November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorininae 417 J - CY e—- C 7 Ly / As) THE FAMILY LITTORINIDAE IN THE INDO-PACIFIC Part I. The Subfamily Littorininae by JOSEPH ROSEWATER Division of Mollusks National Museum of Natural History Washington, D.C. 20560 U.S.A. Abstract The classification of tropical Indo-Pacific Lit- torinidae (Mollusca: Mesogastropoda: Lit- torininae) has been revised. Thirty four Recent and six Tertiary fossil species belonging to six generic groups are described, and/or redescribed, and figured. Complete systematic synonymies are given, together with discussions of relationships, biology and zoogeography. One new subgenus and two new subspecies are described; one new name is assigned. Spawn characteristics of Lit- torinidae are reviewed. A list is given of recog- nized taxa of world-wide Littorinidae belonging to the subfamily Littorininae. Indo-Pacific Littorinidae Representatives of the family Littorinidae are found in most regions of the world occupying habitats from relatively shallow waters below the intertidal zone to situations high above the sea where they may be wet only occasionally by spray. Certain species are unusual among marine gastropods in their ability to survive for long periods without immersion in sea water (Rose- water, 1963a). These species may be considered as living examples of some stages in the invasion of the land by marine snails. The genus Crem- noconchus Blanford, 1869, inhabits fresh water, although it is doubtful that it invaded that habitat from the sea. Although the present paper is main- ly concerned with marine Indo-Pacific Lit- torinidae, the list of recognized taxa on page 423 [05-267] contains the names of all valid species recognized by me in the various generic groups. Family Characters Littorinidae are generalized mesogastropods, and members of the type-genus, Littorina, have nothing outstanding about their appearance. It is, perhaps, this generalized condition which helps to distinguish them from other closely related gastropod families which possess at least some prominent characteristics. “Typical” Littorina, best exemplified by the type-species Littorina littorea (Linné, 1758) (see pl. 326, figs. 1, 2) have thick-walled turbinate shells, are usually non- umbilicate, and have paucispiral opercula, al- though members of the genus Echininus disre- gard the general rule and have umbilicate shells and multispiral opercula. Sculpture varies from nearly smooth to spirally striate, axially furrowed, or nodulose. Some species of Tectarius are quite elaborately sculptured. The anatomy of Littorina was described in some detail by Fretter and Graham (1962, see our pl. 327, figs. A and B). There is a moderately- sized, ditaxic foot bearing the operculum on its dorsal posterior surface; a head with two tentacles having dark eyes at their outer bases; a central, anterior snout with the mouth at its extremity. All species in the family so far as is known are dioecious; the male bears a penis and the female has a well-marked groove on the side of the “neck” for passage of eggs. Fertilization is in- ternal, and pelagic development is the rule, al- though several species have evolved ovovivipari- ty. Species differences may be noted in penile anatomy, reproductive habits and characters of egg capsules (see table). [05-261] 418 Littorinidae Joseph Rosewater Littorinidae ayaa gai Cee y ii}; ; 1 i oii Plate 325. Subfamily Littorininae (explanation on opposite page). (all figures about natural size) [05-262] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorininae 419 Littorinidae most often are confused with such families as Planaxidae. The resemblance between the two is considerable but superficial. This con- fusion may be dispelled if it is remembered that the columella of Planaxidae terminates in a short, “pinched” siphon, while that of Littorinidae is always simple. The closest marine relative, Lacu- nidae, is readily recognized, due to the omnipre- sent umbilical chink, usually more conical shells, and metapodial tentacles. Details of habitat and reproductive characters are also different. Other possible Littorina “look-alikes” may be found among the Fossaridae and Trochacea and also in certain land and freshwater prosobranch families, such as the Helicinidae and Hydrobiidae. The latter do not, of course, occur in a habitat similar to that occupied by Littorinidae. Trochacea differ significantly in having the interior of the shell nacreous and also in being quite different ana- tomically. Throughout the descriptions of the species, measurements of shell size are given in both millimeters and inches. The maximum length to which shells grow is given. The factor of obesity was determined by dividing the width of the shell by the length. Spawn Characteristics A review of the literature on reproduction in Littorinidae reveals that most species, about which the information is known, release pelagic capsules. A few deposit egg masses on a suitable substrate and produce living young. Below is a partial list of species, arranged according to spawn type and faunal region. Following each species I have given the references and, where necessary, the correct name in square brackets. A. Pelagic Capsule— Indo-Pacific: Littorina brevicula: Kojima (1957, 1958a); Habe (1958); Yamamoto and Habe (1962); Hirai (1963); Yamaguchi (1967). L. coccinea (per- sonal observations, see under Description of L. coccinea). L. pintado: Ostergaard (1950); Whip- ple (1965); Struhsaker (1966). L. strigata [= L. undulata?]: Kojima (1958c); L. undulata (per- sonal observations, see Description of L. undulata). Nodilittorina pyramidalis: Kojima (1958b). Gran- ulilittorina granularis [= G. millegrana]: Tokioka and Habe (1953); Habe (1955); Yamamoto and Habe (1962); Hirai (1963). G. picta: Whipple (1965); Struhsaker (1966, 1968b). North Pacific: Littorina squalida: Habe (1958); Kojima (1958b, e); Yamamoto and Habe (1962). Eastern Pacific: Littorina planaxis: MacGinitie and MacGinitie (1949), Gibson (1964). Western and Eastern Atlantic: Littorina lit- torea: Thorson (1946). Eastern Atlantic: Littorina neritoides: Lebour (1935). Western Atlantic: Littorina ziczac: Lebour (1945); Abbott (1954); Lewis (1960); Marcus and Marcus (1963); Borkowski (1969). L. flava: Mar- cus and Marcus (1963). L. lineata: Borkowski (1969). L. lineolata: Borkowski (1969). L. melea- gris: Lewis (1960). Nodilittorina tuberculata: Le- bour (1945); Abbott (1954); Lewis (1960). Tecta- rius muricatus: Lebour (1945); Abbott (1954); Lewis (1960). Explanation to plate 325 (opposite page) Figs. 1-4. Littorina (Littorinopsis) scabra scabra (Linné), showing some of the variation in color and form exhibited by the shell of this species. 1. from Pulau Hantu, SW of Keppel Harbor, Singapore (USNM 660841); 2. from Can- daranan Id., Balabac, Philippines (USNM 233258); 3. from N. Queensland, Australia (USNM 149898); 4. from Bohaydulong Id., North Borneo (USNM 658031). Fig. 5. Littorina (Littorinopsis) carinifera (Menke) from Bombay, India (USNM 90470). Figs. 6, 7. Littorina brevicula (Philippi) from Awaji, Japan (USNM 343538). Figs. 8-10. Littorina (Littoraria) undulata Gray. 8. from Pulau Nias, SW Sumatra, Indonesia (USNM 654451); 9, 10. from N. shore of Guadalcanal Id., Solomon Ids. (USNM 598188). Figs. 11,12. Littorina (Littoraria) coccinea (Gmelin) from SW Viti Levu, Fiji (USNM 531795). Figs. 13,14. Littorina (Littoraria) mauritiana (Lamarck) from Mauritius (USNM 26744), Figs. 15,16. Littorina (Littoraria) pintado pintado (Wood) from Hilo, Hawaii (USNM 339401). Figs. 17,18. Littorina (Austrolittorina) unifasciata unifas- ciata Gray. 17. from Port Jackson, New South Wales (USNM 89472): 18. from Kalbarri, Western Australia (USNM 691672). Figs. 19,20. Nodilittorina australis (Gray). 19. from Rott- nest Id., Western Australia (USNM 671214): 20. from Port Denison, Western Australia (USNM 691677). Figs. 21,22. Nodilittorina nodosa (Gray). 21. from “Western Australia” (USNM 679494): 22. from Kalbarri, Western Australia (USNM 691680). Fig. 23. Littorina (Littoraria) kraussi Rosewater, from Mau- ritius (USNM 89410). Figs. 24,25. Nodilittorina pyramidalis (Quoy and Gaimard) from Lord Howe Id., off New South Wales, Australia (USNM 684715). Figs. 26,27. Littorina (Littoraria) praetermissa May from Port Fairy, Victoria, Australia (USNM 637353). Figs. 28,29. Littorina (Littorinopsis) melanostoma Gray. 28. from Pulau Lumut, Port Swettenham, Malaysia (USNM 661028); 29. from Kranji, Singapore (USNM 631930). [05-263] 420 Littorinidae B. Egg Mass— North and Northeastern Pacific: Littorina at- kana: Kojima (1958a, d); Habe (1958). L. sitkana: Habe (1958); Yamamoto and Habe (1962). Western and Eastern Atlantic: Littorina ob- tusata: Thorson (1946). C. Ovoviviparous— Indo-Pacific: Littorina scabra scabra: Whipple (1965); Struhsaker (1966). Eastern and Western Atlantic: L. scabra angu- lifera: Lebour (1945); Lenderking (1954); Marcus and Marcus (1963). L. saxatilis: Thorson (1946). Joseph Rosewater Littorinidae In general, based on available information, most members of the subgenera Littorina s.s. and Littoraria produce a plano-convex or bi-convex egg capsule, containing from one to a dozen or so eggs (L. littorea, brevicula, squalida, undulata, coccinea, pintado). Information regarding the na- ture of the capsule is lacking for some species, such as L. planaxis. In the cases of L. obtusata, sitkana and atkana an egg mass is deposited on the substrate and, of course, L. saxatilis is ovovi- viparous. Members of the subgenus Littorinopsis for which data are available are also ovovivipa- rous. In Austrolittorina and Nodilittorina rather Plate 326. Type-species of Genera included in Indo-Pacific Littorinidae, Part I. Figs. 1,2. Littorina (Littorina) littorea (Linné, 1758); speci- men figured in J. G. Jeffreys, 1865, British Conchology, Vol. 3, pl. 8, fig 3 (Jeffreys Collection, USNM 185523; 29.7 x 21.3 mm.) Figs. 3,4. Littorina (Littorinopsis) angulifera (Lamarck, 1822) from Sabanilla, Colombia (USNM 103151; 34.1 x 19.1 mm.) Fig. 5. Littorina (Austrolittorina) unifasciata unifasciata (Gray, 1826) from Port Jackson, Australia (USNM 89472; 20.9 x 12 mm.; for abapertural view see pl. 359, fig. 2). Figs. 6,7. Littorina (Littoraria) zebra (Donovan, 1825) from Venado River, Panama Canal Zone (USNM 589696; 29.6 x 19.9 mm.) Figs. 8,13. Nodilittorina (Granulilittorina) millegrana (Phi- lippi, 1848) from Pulau Jerak, West of Sembilan Islands, Malaysia (USNM 661049; 12.1 x 8 mm.). Figs. 9,11. Nodilittorina (Nodilittorina) pyramidalis (Quoy and Gaimard, 1832) from Airport Beach, Barrow Island, Western Australia (USNM 691681; 11.9 x 7.2 mm.) Figs. 10,12. Nodilittorina (Echinolittorina) tuberculata (Menke, 1828) from Permé, northwest of Cape Tiburon, Atlantic coast of Panama (USNM 664217; 16.4 x 11 mm.) [included for comparison, although there are no Indo- Specific species]. [05-264] November 30, 1970 elaborate ““gear-decorated” or “‘terraced” capsules are produced. Of the three species whose egg capsules are known in Melarhaphe, one produces a flattened cylinder-shaped capsule (L.neritoides) another a tiered capsule (L. meleagris) and the third a biconvex capsule (L. flava) [the generic assignments of the last two forms is still question- able]. Tectarius muricatus produces a capsule which is rather like that of L. neritoides, but is biconvex. A definite systematic trend in the shapes of littorinid egg capsules does not seem to exist Plate 327. Littorina littorea (Linné) removed from shell show- ing right side of body of A. female; B. male (from Fretter and Graham, 1962). Abbre viations— INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorininae 421 ag, albumen gland OV, ovary em, columellar muscle p, penis cov, covering gland png, penial glands cpg, capsule gland pr, prostate dg, digestive gland ro, renal section of oviduct e, eye sf, sole of foot ie foot sgr, sperm groove hg, hypobranchial gland © sn, snout k, kidney sts stomach me, mantle edge ts tentacle op, operculum td, testicular duct acting as opr, ovipositor vesicula seminalis above the species level. It does appear, however, that each species has fairly characteristic spawn. Radulae The radulae of Littorinidae may be described as generalized taenioglossate (see pl. 328). So far as is known most species browse on the epiphytes of the hard substrates upon which they live. The radulae are adapted for scraping off this food and may become very long as considerable replace- ment of worn teeth is required. The unused portion is carried coiled up in the radula sac behind the head. Radulae of Littorina and its subgenera are all fairly similar, having the formu- la 2-1-1-1-2, and consisting of a slender multicus- pid outer marginal, (pl. 328 D) a more robust inner marginal (pl. 328 C), a subquadrate lateral with a basal embayment (the embayment is char- acteristic of Littorinidae ) (pl. 328B), and a rachi- dian or central which is typically about as high as it is wide and which bears 3 cusps. In Nodilit- torina and Tectarius there is a tendency for narrowing of the central tooth although the lateral and marginals are quite similar to the condition in Littorina. In the genus Echininus and especially its subgenus Tectininus, as shown by Abbott (1954), there is considerable reduction in width of both central and lateral teeth. Since most mem- bers of the Littorinidae apparently share the gen- eralized taenioglossate radula and squarish lateral teeth, it does not seem to offer much promise for systematic diagnosis below the generic level. Distribution Most Littorinidae inhabit tropical seas, and the largest numbers of species live in the Indo- Pacific region. A conservative estimate places the total number of Recent world-wide Littorinidae of all genera at slightly above 100 species, exclud- ing the freshwater representatives which at least may be considered a separate subfamily of the Littorinacea as may the members of the Rissel- (UA) PAA? 7 7 F TO Da ee a os L_Olmm__ 5 Plate 328. Radula of Littorina littorea (Linneé), type-species of the genus Littorina. One complete transverse row of many which make up radula ribbon. Fig. A. rachidian or central tooth; Fig. B. lateral tooth C. and D. inner and outer marginal teeth. [05-265 ] 422 Littorinidae Joseph Rosewater Littorinidae lidae. Of these, some 34 species are to be found in the tropical Indo-Pacific, 21 inhabit the eastern Pacific, 16 the western Atlantic, only 9 the eastern Atlantic (5 of which are shared with the western Atlantic) and 27 more or less have been enumerat- ed in the South Ocean (Antarctic) (see List of Recognized Taxa). Representatives of the family are conspicuously absent from the southern tip of South America. The distributional pattern is further complicated by the present in two or more regions of what are termed analogues, species which bear considerable resemblance and may be related (Rosewater, 1963b). Since the geologic history of the Littorinidae dates from the Jurassic, it is doubtful that origins of the family can be unraveled. Dispersal of species is aided now and undoubtedly was helped in the past by pelagic larvae and the semi-arboreal habitat of such species as L. scabra. A single fertilized female of the latter oviparous species, if rafted to a favor- able locality, could establish a new colony. Classification Adapted as they are for intertidal life, it is not surprising to find little variation in general pat- tern of littorinid speciation. The shells are usually rounded-turbinate or some modification of this shape, permitting the animals to cling to rocks in surf or creep into crevices to avoid dessication. Grossly, the animals which form the shells do not vary much either. The systematists’ task of classi- fying species and groups of species into genera has come to depend on consistent differences in shape and sculpture and outstanding differences in anatomical details. Characters which have been utilized most in the present study are ab- sence or presence of nodulosity and granulate sculpture, overall color pattern and shape, the radula, and penile anatomy. The latter character appears to be an extremely important and reliable one, and two levels of classification have been based on it, at least in part: 1) the separation of species and 2) the formation of species groups (genera) exhibiting overall similarity in penile anatomy and other features. Difficulties have been encountered in the clas- sification of fossils. As mentioned previously, shells of Littorinidae resemble those of a number of other marine, freshwater and land _proso- branchs. If fossil preservation is not good, the decision on familial placement is extremely dif- ficult, if not impossible. Even if preservation is optimum, since there are no anatomical charac- ters upon which to rely, one can almost never be sure that these assignments are correct. This is especially true of the older fossils where resem- blance to Littorinidae is rather nebulous. For these reasons much concerning fossils has been taken on faith and must be regarded in that light. Only Tertiary Indo-Pacific fossils are included here or listed in the List of Recognized Taxa. As is the case with many groups of animals, not alone mollusks, an ample number of species and genera have been described in the Littorinidae. In many cases these are not justified in the light of modern systematics and some have had to be synonymized. Early in the course of this study it was thought probable that most of the generic names erected for species originally described or early assigned to Littorina would be of little value. After long evaluation, it has become ap- parent that some of these names may have real utility, at least as subgeneric groups indicators and it may be noted that they are so used through- out. It is hoped that in each case the reason for their use will be made clear. List of Recognized Taxa The following is a list of recognized species, subgenera and genera of world Littorinidae be- longing to the subfamily Littorininae. Fossils listed are Tertiary forms described from the Indo- Pacific region alone; the sheer numbers and the uncertainties surrounding extraterritorial and pre-Tertiary fossil species preclude their con- sideration here. As mentioned previously, the generic groupings are controversial but are of- fered as a working solution to some method of orderly arrangement of species. Due to the means of presentation many possibly familiar names will be missed by some in the list. Often they are considered to be synonyms and in the cases of Indo-Pacific species are included in the synony- mies which follow. The few fossil species names are preceded by a dagger [f]. Certain littorinoid groups, although definitely “Indo-Pacific” in general geography, are not con- sidered here. These are the members of such genera as Bembicium and Peasiella here believed to constitute at least a separate subfamily of Littorinidae. I consider the freshwater forms, such as Cremnoconchus and Cremnobates, also to be separable as at least a subfamily. Forms having definite southern ocean affinities, inhabiting New Zealand and the Antarctic, such as Laevilitorina, Macquariella, etc., are included in the list of recognized taxa, but will not be considered in detail here. A second part of Indo-Pacific Lit- torinidae, covering the subfamilies Tectariinae and Echininae will appear at some future time. [05-266] November 30, 1970 Family Littorinidae Gray, 1840 Subfamily Littorininae GENUS Littorina Ferussac, 1822 Subgenus Littorina Ferussac, 1822. littorea (Linné, 1758). Type. Recent, North Atlantic obtusata (Linné, 1758). Recent, North Atlan- tic saxatilis (Olivi, 1792). Recent, circumboreal sitkana Philippi, 1846. Recent, northeast Pacific atkana Dall, 1886. Recent, North Pacific aleutica Dall, 1872. Recent, North Pacific squalida Broderip & Sowerby, 1829. Recent, North Pacific brevicula (Philippi, 1844). Recent; East Asia, Japan planaxis Philippi, 1847, Recent, northeast Pacific scutulata Gould, 1849. Recent, northeast Pacific ziczac (Gmelin, 1791). Recent; tropical wes- tern Atlantic Subgenus Littoraria Gray, 1834 zebra (Donovan, 1825). Type. Recent, tropi- cal eastern Pacific undulata Gray, 1839. Recent, Indo-Pacific nebulosa (Lamarck, 1822). Recent, Carib- bean and Gulf of Mexico mauritiana (Lamarck, 1822). Recent, Indian Ocean kraussi Rosewater, new name Recent, Indian Ocean coccinea (Gmelin, 1791) Recent; Indo- Pacific praetermissa May, 1909. Recent, South Aus- tralia pintado (Wood, 1828). Recent; Indo-Pacific pintado schmitti, Bartsch & Rehder, 1939. Recent, Clipperton Island pullata Carpenter, 1864. Recent; tropical eastern Pacific cingulifera Dunker 1845. Recent; tropical west Africa ?sundaica Altena, 1945. Recent, Java Pacutispira E. A. Smith, 1892. Recent, Aus- tralia Pinfans E. A. Smith, 1892. Recent, Australia tkozaiensis Nomura and Onisi, 1940. Mi- ocene, Japan tadonis Yokoyama, 1927. Pliocene, Japan tlucida Yokoyama, 1927. Pliocene, Japan INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorininae 423 Subgenus Littorinopsis Mérch, 1876 scabra scabra (Linne, 1758). Recent, Indo- Pacific scabra angulifera (Lamarck, 1822). Type. Recent, eastern and western Atlantic scabra aberrans Philippi, 1846. Recent, trop- ical east Pacific melanostoma Gray, 1839. Recent, south Asia. carinifera (Menke, 1830). Recent, Indo- Pacific irrorata (Say, 1822). Recent, western Atlan- tic fasciata Gray, 1839. Recent, tropical east Pacific varia Sowerby, 1832. Recent, tropical east Pacific modesta Philippi, 1846. Recent, tropical east Pacific paytensis Philippi, 1847. Recent, tropical east Pacific + miodelicatula Oyama, 1950. Miocene, Ja- pan. tincisa Yokoyama, 1927. Pliocene, Japan. Subgenus Austrolittorina Rosewater, new sub- genus unifasciata unifasciata Gray, 1826. Type. Recent, Australia unifasciata antipoda Philippi, 1847. Recent, New Zealand unifasciata fernandezensis Rosewater new subspecies. Recent, Islas Juan Fernandez cincta Quoy and Gaimard, 1832. Recent, New Zealand araucana Orbigny, 1840. Recent, Chili and Peru peruviana (Lamarck, 1822). Recent, south- eastern Pacific aspera Philippi, 1846. Recent, tropical east Pacific penicillata Carpenter, 1864. Recent, Baja California tessellata Philippi, 1847. Recent, Caribbean lineata Orbigny, 1841. Recent, tropical wes- tern Atlantic lineolata Orbigny, 1840. Recent, tropical western Atlantic punctata (Gmelin, 1791). Recent, East At- lantic and South Africa africana Philippi, 1847. Recent, South Afri- ca knysnaensis Philippi, 1847. Recent, South Africa 424 Littorinidae Subgenus Melarhaphe Menke, 1828 neritoides (Linne, 1758) Type. Recent, east Atlantic ?mespillum (Muhlfeld, 1824). Recent, tropi- cal western Atlantic ?meleagris (Potiez and Michaud, 1838) Re- cent, West Indies ?umbilicata Orbigny, 1840. Recent, Chili and Peru ? flava King and Broderip, 1832. Recent, tropi- cal west Atlantic Subgenus Algamorda Dall, 1918 newcombiana (Hemphill, 1877). Type. Re- cent, northwest North America GENUS Nodilittorina von Martens, 1897 Subgenus Nodilittorina von Martens, 1897 pyramidalis pyramidalis (Quoy and Gai- mard, 1833). Type. Recent, Indo-Pacific. pyramidalis pascua Rosewater, new subspe- cies. Recent, Easter and Pitcairn Islands. natalensis (Philippi, 1847). Recent, East Af- rica and Madagascar. australis (Gray, 1826). Recent, W. Australia. nodosa (Gray, 1839). Recent W. Australia. galapagiensis (Stearns, 1892). Recent, Gala- pagos Islands. Subgenus Echinolittorina Habe, 1956 tuberculata (Menke, 1828). Type. Recent, tropical west Atlantic. Subgenus Granulilittorina Habe and Kosuge, 1966 millegrana (Philippi, 1948). Type. Recent, Indo-Pacific. subnodosa (Philippi, 1847). Recent, Red Sea and Persian Gulf. leucosticta leucosticta (Philippi, 1847). Re- cent, India. leucosticta biangulata (von Martens, 1897). Recent, east Indies and western Melane- sia. leucosticta feejeensis (Reeve, 1857). Recent, western Pacific. exigua (Dunker, 1860). Recent, Japan. tiwakiana (Nomura and Hatai, 1936). Mi- ocene, Japan. cinerea (Pease, 1869). Recent, Marquesas Islands picta (Philippi, 1846). Recent, Hawaii. Joseph Rosewater Littorinidae ?miliaris (Quoy and Gaimard, 1833). Recent, east Atlantic ?striata (King and Broderip, 1832). Recent, east Atlantic GENUS Laevilitorina Pfeffer, 1886 Subgenus Laevilitorina Pfeffer, 1886 caliginosa caliginosa (Gould, 1849). Type. Recent, Southern Ocean caliginosa aestualis Strebel, 1908. Recent, Antarctica antipoda (Filhol, 1880) Recent, New Zea- land bifasciata Suter, 1913. Recent, New Zealand bennetti Preston, 1912. Recent, Antarctic claviformis Preston, 1916. Recent, Antarctic granum Pfeffer in von Martens & Pfefter, 1886. Recent, Antarctic latior Preston, 1912. Recent, Antarctic pygmaea Pfefter, 1886. Recent, Antarctic umbilicata Pfeffer, 1886. Recent, Antarctic venusta Pfeffer, 1886. Recent, Antarctic antarctica E. A. Smith, 1902. Recent, Ant- arctic Subgenus Corneolitorina Powell, 1951 coriacea (Melvill and Standen, 1907). Type. Recent, Antarctic elongata Pelseneer, 1903. Recent, Antarctic heardensis Dell, 1964. Recent, Antarctic GENUS Macquariella Finlay, 1926 hamiltoni (Smith, 1898). Type. Recent, New Zealand aucklandica Powell, Zealand delli Powell, 1955. Recent, New Zealand macphersonae Dell, 1964. Recent, Mac- quarie Island 1933. Recent, New GENUS Laevilacunaria Powell, 1951 Subgenus Laevilacunaria Powell, 1951 bransfieldensis (Preston, 1916). Type. Re- cent, Antarctic antarctica (von Martens, 1885). Recent, Ant- arctic pumilio (E. A. Smith, 1879). Recent, Antarc- tic Subgenus Pellilacunella Powell, 1951 bennetti (Preston, 1916). Type. Recent, Ant- arctic [05-268] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorininae 425 GENUS Pellilitorina Pfeffer, 1886 setosa (Smith, 1875). Type. Recent, Antarc- tic pellita (von Martens, 1885) Recent, Antarc- tic GENUS Rissolittorina Ponder, 1966 alta (Powell, 1940). Type. Recent, New Zea- land mariae (Tenison-Woods, 1876). Recent, Tas- mania Abbreviations The following institutional abbreviations are used in this paper: AMS-Australian Museum, Sydney ANSP-Academy of Natural Sciences of Phil- adelphia BM(NH)-British Museum (Natural History}, Lon- don BPBM-Bernice P. Bishop Museum, Honolulu MHNG-Muséum d Histoire Naturelle, Geneva MHNP-Muséum d'Histoire Naturelle, Paris MCZ-Museum of Comparative Zodlogy, Cam- bridge, Massachusetts NMW-National Museum of Wales, Cardiff RNHL-Rijksmuseum van Natuurlijke Historie, Leiden SMF-Senckenberg Museum, Frankfurt-am-Main USNM-United States National Museum, Wash- ington, D.C. WAM-Western Australian Museum, Perth, West- ern Australia YPM-Peabody Museum of Natural History, Yale University, New Haven, Connecticut ZMA-Zoological Museum, Amsterdam ZMC-Zoological Museum, Copenhagen Acknowledgements The following persons contributed to making this study more complete by providing informa- tion, research assistance, specimens or working space; their help is gratefully acknowledged: R. T. Abbott, Delaware Museum of Natural History; C. O. van Regteren Altena, E. Git- tenberger, R. M. van Urk-RNHL; E. Binder- MHNG; J. B. Burch-Museum of Zoology, Uni- versity of Michigan; W. J. Clench, R. D. Turner, kK. J. Boss-MCZ; H. Coomans-ZMaA; S. P. Dance- NMW; R. K. Dell-Dominion Museum, Wel- lington, New Zealand; E. Fisher-MHNP; L. Fish- elson-Tel-Aviv University, Israel; W. D. Hart- man-YPM; Johnny Johnson and Mr. and Mrs. D. Everett-Singapore; A. M. Keen-Stanford Univers- ity; J. Knudsen, H. Lemche-ZMC; Y. Kondo- BPBM; S. Kosuge-National Science Museum, Tokyo; H. S. Ladd-U. S. Geological Survey; D. F. McMichael-AMS; R. Robertson and V. Orr Maes- ANSP; H. Steinitz-Hebrew University, Jerusa- lem, Israel; Jeannette Whipple Struhsaker- University of Hawaii, Honolulu; N. Tebble, J. F. Peake-BM(NH); G. Thorson-Marine Biological Lab, Helsinggr, Denmark; B. R. Wilson-WAM; A. Zilch-SMF; Mrs. Ann Cohen called to my atten- tion specimens she collected in the Juan Fer- nandez Islands; E. Brody, A. Griffin, J. Green- field, R. Capen, Research Assistants; H. A. Rehder, J. P. E. Morrison, C. F. E. Roper, Divi- sion of Mollusks, USNM. The U.S. Atomic Energy Commission, through I. E. Wallen and R. W. Hiatt, arranged for me to work at the Eniwetok Marine Biological Labora- tory. A portion of this work was supported by the National Science Foundation as a part of the U.S. Program in Biology, International Indian Ocean Expedition. The work was also supported in part by Smithsonian Institution Research Foundation Grant number Sg 0684025. Doubtful Species of Littorinidae Included under this heading are species of Littorinidae from the Indo-Pacific which I con- sider either unrecognizable as valid species from the faunal area, or as being questionably included in the family. In the latter category, some species already have been removed from the family by prior workers. Although Iredale and McMichael (1962) listed Larinopsis turbinata (Gatliff and Gabriel, 1909) and L. ostensus Iredale, 1936, in Littorinidae, the genus Larinopsis Gatliff and Gabriel, 1916, was placed in Fossaridae by Wenz (1940) and by Macpherson and Gabriel (1962, including the mentioned species). Ponder (1966) has shown that some species placed in Zelaxitas, formerly thought to be Littorinidae, belong to three different families, that genus now being assigned to Rissoellidae. Only one species origi- nally included under Zelaxitas, Z. alta Powell, 1940, remains in Littorinidae, although in a dif- ferent genus (see List of Recognized Taxa under Rissolittorina Ponder). Littorina beccarii Tapparone-Canefri, 1875 Range—Known only from the type-locality: So- rong, northwest New Guinea (West Irian, Indone- sia); [05-269] 426 Littorinidae Remarks—This species has not been figured and its identity is in doubt. The author’s sugges- tion that an affinity exists with Littorina lamellosa Montrouzier indicates that it is a Fossarus and not a Littorina. Mrs. van Benthem Jutting (1962) published the results of her studies on the types of New Guinea non-marine mollusks described by Tapparone-Canefri. These were preserved in the Genoa Museum, and it is possible that the type of L. beccarii is there also. Unfortunately it has not been possible to ascertain this fact in the course of the present study Synonymy— 1875 Littorina beccarii Tapparone-Canefri, Annali del Museo civico di Storia Naturale di Genova, vol. 7, p. 1031 (Sorong [N.W. New Guinea]). Littorina fragilis Fenaux, 1943 Remarks—The illustration accompanying the description of this species indicates that it is very probably a member of the genus Diala (Ce- rithiidae) and not a Littorina. Synonymy— 1943 Littorina fragilis Fenaux, Bulletin de l'Institut océan- ographique, Monaco, no. 835, p. 7, figs. 4,5 (Pau- motou). Problitora globulus (Angas, 1880) Remarks—Macpherson and Gabriel (1962) placed this species in the genus Problitora Ire- dale, 1931, of which P. moerchi (Adams and Angas) is the type-species. As globulus was de- scribed from shell-sand, and has been collected on only very few occasions, its true relationships remain to be proven. At present there appears to be little reason for retaining these peculiar forms in the family Littorinidae as they resemble much more closely members of the family Naticidae. Synonymy— 1880 Amauropsis globulus Angas, Proceedings of the Zoologi- eal Society of London, 1880, part 3, p. 416, pl. 40, fig. 5 (Holdfast Bay in shell-sand [near Adelaide, South Australia]); types, BM(NH) 81.4.29.5. Littorina hisseyiana Tenison-Woods, 1876 Remarks—May (1903) figured the type of L. hisseyiana. His illustration shows a tiny, ap- parently umbilicated snail with a multispiral operculum, and having a color pattern resembling Joseph Rosewater Littorinidae one of the Trochacea. It is certainly not a Lit- torina. Synonymy— 1876 Littorina hisseyiana Tenison-Woods, Papers and Pro- ceedings of the Royal Society of Tasmania, for 1875, p. 148 (from stomach of a mullet, Agenostoma die- menensis Rich, probably caught in the Derwent [Tas- mania]). 1903, W. L. May, ibid., for 1902, p. 111, fig. 7; 7 syntypes in Tasmanian Museum (T.M. 5480) according to Hardy, 1916, ibid., for 1915, p. 66. Littorina lamellosa Montrouzier, 1861 Remarks—Although described as a Littorina, the author observed that this species resembles a fossarid. This is certainly the case, as the original illustration almost exactly matches the Hawaiian species which was named by Pease as Fossarus garretti. Synonymy— 1861 Littorina lamellosa Montrouzier, Journal de Conchyliol- ogie, vol. 9, p. 273, pl. 11, fig. 5 ({Ile] Art, New Caledonia). Problitora moerchi (Adams and Angas, 1864) Remarks—See Remarks under P. globulus (An- gas). I consider this species not to belong in Littorinidae. The figures by Hedley (1902) es- pecially that of the operculum, resemble more closely one of the Naticidae. Synonymy— 1864 Amauropsis moerchi Adams and Angas, Proceedings of the Zoological Society of London, 1863, part 3, p. 423 (Watson’s Bay, Port Jackson [New South Wales]); type, BM(NH) 70.10.26.178. 1902, Hedley, Proceed- ings of the Linnean Society of New South Wales, vol. 26, part 4, p. 700, pl. 34, figs. 19, 20. Littorina reticulata Anton, 1839 Remarks—Anton’s species could be any of sev- eral Granulilittorina. The description and Philip- pis figure are too vague to identify the species with any precision and the lack of a locality increases the likelihood that L. reticulata may be an extra-Indo-Pacific entity. The species is here considered unrecognizable. Synonymy— 1839 Litorina reticulata Anton, Verzeichniss der Conchylien welche sich in der Sammlung von H. E. Anton befinden. Halle, p. 53 (no locality given). 1847, Phi- lippi, Abbildungen und Beschreibungen Conchylien, vol. 2, p. 199, pl. 4, fig. 12 [figure said to be drawn from one of Anton’s type-specimens]. 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Publications of the Seto Marine Biological Laboratory, vol. 1, no. 3, pp. 151-152, fig. Tokioka, T. and T. Habe. 1953. A new type of Littorina- capsula. ibid., vol. 3, no. 1, pp. 55, 56, fig. Tryon, G. W. 1887. Family Littorinidae. Manual of Concholo- gy, vol. 9, pp. 229-313, pls. 40-51. Von Martens, E. 1887. List of the shells of Mergui and its Archipelago, collected for the Trustees of the Indian Muse- um, Calcutta by Dr. John Anderson, F.R.S., Superintendent of the Museum. The Journal of the Linnean Society, Zo- ology, vol. 26, no. 130, pp. 155-219, 3 pls. Weinkauff, H. C. 1856-1882. Die Gattung Litorina. Systemat- isches Conchylien-Cabinet, edition 2, vol. 2, prt. 9, pp. 1-114, 14 pls. Weinkauff, H. C. 1883. Catalog der Gattung Litorina Feru- ssac. Jahrbuch der Deutschen Malakozoologischen Gesell- schaft, vol. 10, pp. 213-227. Wenz, W. 1940. Handbuch der palaozoologie, vol. 6, prt. 1 (6), pp. 721-960, figs. 2084-2787. Whipple, Jeanette A. 1965. Systematics of the Hawaiian Littorina Férussac (Mollusca: Gastropoda). The Veliger, vol. 7, no. 3, pp. 155-166, pls. 25, 26, 4 text figs. Whipple, J. A. [is maiden name of J. W. Struhsaker, q.v.]. Winckworth, R. 1922. Nomenclature of British Littorinidae. Proceedings of the Malacological Society of London, vol. 15, prts. 2, 3, pp. 95-97. Yamaguchi, M. 1967. Egg capsules of a Periwinkle, Littorina brevicula, in plankton samples. The Venus, vol. 25, no. 2, pp. 73-76, figs. 1, 2. Yamamoto, G. and T. Habe. 1962. Fauna of shell-bearing mollusks in Mutsu Bay. Scaphopoda and Gastropoda (1). Bulletin of the Marine Biological Station of Asamushi, Tohoku University, vol. 11, no. 1, pp. 1-20, pls. 1-3. Yen, Teng-Chien. 1942. A review of Chinese Gastropods in the British Museum. Proceedings of the Malacological Society of London, vol. 24, parts 5, 6, pp. 170-289, pls. 11-28. [05-277] 430 Littorinidae Joseph Rosewater Littorinidae Subfamily Littorininae Gray, 1840 Key To Subgeneric Groups of Indo-Pacific Littorininae The following key is an attempt to provide a point of departure for the placement of a littorinid species in a particular subgeneric group. Even at the generic level there is a great deal of phe- notypic variation expressed and it is often true that such hopefully dependable characters as nodular and granular sculpture are evanescent. Other keys are provided in the appropriate places to species of Littoraria and Littorinopsis. None are provided for the other groups, however, be- cause it was felt that species of Austrolittorina and Nodilittorina s.s. are sufficiently distinct and geographically situated so as to essentially pre- clude difficulties in identification. In the case of Granulilittorina, the nature of speciation in the group makes the construction of an effective key a near impossibility. It is suggested that the user may wish to acquire some familiarity with the range of variation in this subfamily before pro- ceeding. 1. Shell nodulose, granulose, or with surface axially folded ......... 0.00.00 ceca 2 1. Shell surface smooth or spirally folded (carinate) .. 0.0.0.0... 00 eee ee ee ee 2. Sculpture nodulose, limited to 2-3 rows per whorl of major nodules, or surface axially folded ee ee ee ee Nodilittorina p. 05-375 2. Sculpture granulose, often in excess of 3 rows of granulations, sometimes nearly smooth lite Bn BuO oniak te a eg 4A ace, BAe Op ats Be ies Soles Granulilittorina p. 05-395 3. Base of shell adjacent to columella flattened or hollowed out, forming a crescent-shaped BIS co deashs Cale Bee oe SE Rae Sa eee ea ee Ae Owes Austrolittorina p. 05-351 3. Base of shell adjacent to columella generally smoothly rounded, not flattened OO SC ONDE - seid hh a ae Ge eek Gods bee bs oe ea OEE BESS SRB OSu Cte ESOG 4 4. Spire less than half the length of shell, multiple penial glands present ahed Bad Se Sea Gee Bee ws dG ee a oa wee wees Littorina p. 05-285 4, Spire usually more than half the length of shell, penial glands absent ...........5 5. Shell rather thin for its size, conspicuously colored or patterned, usually arboreal. or plami-living Species: fei. avulk eee be Dene BS Littorinopsis p. 05-329 5. Shell not thin for its size, often with subdued coloration or lacking pattern, usually roclk-livine Species’ 24 dcn cdo beee nd deena ee oS ea Littoraria p. 05-297 November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 431 Family Littorinidae Gray, 1840 Subfamily Littorininae Gray, 1840 Genus Littorina Férussac, 1822 Type: Littorina littorea (Linneé, 1758) Considered broadly, the genus Littorinaencom- passes a multitude of Recent and fossil species throughout the world. The impossibility of prop- erly classifying the large number of poorly pre- served fossil species described from the Upper Paleozoic through the Tertiary prevents coverage here of any but Tertiary Indo-Pacific fossils. Living species of the genus Littorina usually occupy habitats on rocks in the intertidal zone although some species prefer shore vegetation. The geographic range of the genus is from Arctic to Antarctic shores, including intervening tem- perate and tropical areas where a suitable habitat is available. The subgenus Littorina, sensu stricto, however, is a northern group and there is only one species of that subgenus whose range extends into the tropical Indo-Pacific, Littorina brevicula (Philippi), which is treated here (see List of Recognized Taxa). Synonymy— 1822 Littorina Ferussac, Tableaux Systematiques des Ani- maux Mollusques, p. xxxiv. Type-species by subse- quent designation, Blainville, 1828, Dictionnaire des Sciences Naturelles, Paris, vol. 56, p. 98: Turbo littoreus [= Littorina littorea (Linne)]. 1827 Littorina “Ferussac’ Nilsson, Petrificata Suecana Forma- tionis Cretaceae, part 2, Mollusca, p. 11. Invalid type-species by monotypy: Turbo sulcatus Nilsson, 1827, a species not originally included by Feérussac. 1827 Neritoides T. Brown, Illustrations of the Conchology of Great Britain and Ireland, Index p. iv, pl. 43, figs. 14, 15, 21, 22. Type-species by monotypy: Neritoides littoralis [=Littorina obtusata (Linne, 1758)]. 1828 Litorina Menke, Synopsis Methodica Molluscorum, p. 24 [emendation of Littorina Ferussac, 1822]. 1847 Littorelaea Leach, Annals and Magazine of Natural History, vol. 20, p. 271; in combination Littorelaea pultneyii [nomen nudum|. 1849 Litormia Morch, Indbydelsesskrift til de oftentlige af- gangs-og aarsprgver i Nykjdbing katedralskole, p. 63 {error for Littorina Feérussac, 1822]. 1854 Bacalia H. and A. Adams, Genera of Recent and Fossil Mollusks, vol. 1, p.312; based on Bacalia Gray, 1840, Synopsis of the Contents of the British Museum, ed. 42, p. 147 [nomen nudum]; type-species by subse- quent designation, Winckworth, 1922: Turbo lit- toreus Linne. 1861 Littorrina Gabb, Proceedings of the American Philo- sophical Society, vol. 8, p. 114 [error for Littorina Férussac, 1822 1869 Neritrema Récluz, Actes de la Société Linnéenne de Bordeaux, series 3, vol. 7 (vol. 27) pp. 43, 46. Type- species by subsequent designation, Dall, 1909, U.S. Geological Survey Professional Paper 59, p. 79: Li- ttorina obtusata Linne,. 1878 Litonia Weinkauff, Systematisches Conchylien-Cabinet, vol. 2, part 9, p. 40 [error for Littorina Ferussac, 1822] 1884 Litorinna Dall, Proceedings U.S. National Museum, vol. 7, p. 344 [error for Littorina Ferussac, 1822] 1891 Isonema ‘Hall’ Provancher, Faune Canadienne; Les Mollusques de la Province de Quebec, part 1, p. 91. Type-species by subsequent designation Bequaert, 1943 Johnsonia, vol. 1, p. 1: Turbo littoreus Linne, 1758; not Isonema Meek and Worthern, 1866. 1891 Littornia Jenkins and Grocock, Hardwicke’s Science Gossip, vol. 27, p. 9; 1917, H. B. Preston, Zoological Record, vol. 52, section 8, Mollusca, p. 33 [error for Littorina Férussac, 1822]. 1918 Algaroda Dall, Proceedings of the Biological Society of Washington, vol. 31, p. 137; type-species by original designation: Littorina littorea Linne. 1918 Littorivaga Dall, ibid., Type-species by original designa- tion: Littorina sitchana Philippi. 1939 Neritotrema ‘Recluz’ Wenz, Handbuch der Palaozoo- logie, Lief 4, Band 6, Gastropoda, part 3, Proso- branchia p. 518 [emendation of Neritrema Récluz, 1869]. 1958 Exzolittorina Habe, Publications from the Akkeshi Ma- rine Biological Station, no. 8, p. 9; type-species by monotypy: Ezolittorina squalida Broderip and Sow- erby. As has been accurately pointed out by Winck- worth (1922) and Bequaert (1943) Férussac cited the names of several species in association with his description of Littorina. The “Paludina ma- rine’ listed on pages 9-11 of the introduction to Férussac’s work are obtusatus, neritoides, Lit- toreus, muricatus and afer. Following the list Férussac mentioned (p. 11) in vernacular “‘lit- torine’ as a subgenus to contain some species previously assigned to Turbo and Trochus. In the following paragraph he assigned Turbo neritoides of Linné to this subgenus, at least in part. Since the latter cannot be construed as a type designa- tion, it is necessary to seek a later one. Blainville (1824, Dictionnaire des Sciences Naturelles, Par- is, vol. 32, p. 226) gave as an example of Férus- sac’s “Littorine’: Turbo littoralis, referring to Chemnitz vol. 5, t. 185, fig. 1852, nos. 1-18 [1-8] [figures are L. littorea]. This is an invalid subse- quent designation (I.C.Z.N. Art. 69 (a) (i) and (iv)), and the same is true in the case of Pay- raudeau's (1826) description of L. basterotii [=L. neritoides (Linné)]| in association with Littorina. The first valid designation is that of Blainville, (1828 ibid.) who clearly indicated Turbo littoreus, one of the originally included species as the type of the genus. [05-279] 432 Littorinidae Joseph Rosewater [ These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence. | [05-280] Littorinidae November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 433 Subgenus Littorina sensu stricto Characteristics of species belonging to the sub- genus Littorina s.s. include a generally boreal distribution, relatively large, somber colored shells (with a number of exceptions) and the morphological character of the presence in the male of a series of penial glands with accessory flagella on the penis. Littorina brevicula (Philippi, 1844) (Pls. 325, 329, 330) Range—Northern China, Korea and Japan (and boreal Pacific). Remarks—Littorina (Littorina) brevicula is an inhabitant of the boreal Pacific. Although strictly a cool water species, it is included here with species occuring in the tropical Indo-Pacific be- cause its range does extend to the southern most of Japan’s main islands and also to the coast of China at latitudes roughly on a level with Tai- wan. It occurs therefore within the range of a number of tropical species. The anatomy of the penis of L. brevicula, which possesses multiple penial glands, the char- acteristics of the floating egg (helmet) capsule, which is a relatively simple plano-convex shape, the generalized radula and somber coloration of the shell all align this species with the boreal group of Littorina s.s. Habitat—Intertidal on rocky coasts. Plate 329. Littorina brevicula (Philippi). Figs. 1,2. Specimens from Takami, Honshu, Japan (USNM 424296; 11.7 x 9.3 mm.) Figs. 3,4. Littorina souverbiana Crosse, Holotype (Journal de Conchyliologie Collection, Paris Museum; 11.5 x 11 mm.) Figs. 5,6. Turbo (Litorina) breviculus Philippi, lectotype figure (from’ Abbildungen und Beschreibungen Con- chylien, vol. 2, p. 161, pl. 3, fig 10; about 11 x 10 mm.) Figs. 7,8. Littorina balteata Reeve, lectotype (British Muse- um (NH) 1968316; 11 x 7.6 mm.). Description—Shell reaching nearly 23 mm. (about 0.8 inch) in length, broadly-to depressed turbinate in shape, often appearing almost as broad as long, but having an average obesity of about .74 (39 specimens range from .68-.79): rather thick in structure, imperforate; usually displaying an accentuatedly pointed siphonal trough; body whorl often sculptured with 3 or 4 raised strong, white-speckled spiral cords. Ex- ternal color variable; reddish brown to grayish white solid color or with banding, spotting, strip- ing and zigzag color patterns. Aperture diffuse tan to brown with faint narrow white band revolving inward from near anterior junction of outer lip and columella, occasionally a wider, very diffuse band visible inside outer lip midway to its anteri- or junction with columella; columella grayish white anteriorly, merging with aperture color posteriorly; inner edge of outer lip often dark brown or speckled brown and white. Base dis- tinctly flattened with comparatively low spiral cords; whorls broadly shouldered below suture. Whorls 4-5, rounded, nuclear whorls usually e- roded. Spire usually less than half the length of shell, produced at an angle varying from about 75° to 110° in very shortspired individuals. Aper- ture rounded-oval; inner and outer lips relatively thickly produced; with a distinctly pointed si- phonal trough at anterior junction of outer lip and columella, often tinged with brown in young individuals; base adjacent to columella seldom depressed, but occasionally an anomalous umbil- ical chink formed. Suture impressed. Spiral sculpture usually consisting of from 3-4 major raised cords on upper half of body whorl; from Plate 330. Littorina brevicula (Philippi). Fig. A. Radula, showing central and one-half row of teeth (upper scale is 0.125 mm.); Hakodate, Japan. Fig. B. Penis of young specimen from Wuchiuhsu, China (lower scale is 1 mm.) [05-285] 434 Littorina Littorinidae Joseph Rosewater 4-6 minor cords on lower half on base; spire sculpture usually at least partially obliterated. Nuclear whorls about 2-2!/2, smooth, light- brown; post nuclear whorls rapidly becoming spirally sculptured with 1-2 cords. Overall micro- scopic sculpture consisting of fine wavy spiral threads especially noticeable between major cords, and fine, rather regular, oblique axial growth markings. Operculum roundly oval, pau- cispiral, the nucleus wen. Peas and nearly central. Radula littorinoid, 2-1-1-1-2, the central tooth broad. Animal darkly pigmented on surfaces of tenta- cles, snout and foot; the penis of small individu- als (shell length 5-6 mm.) bearing 2 or 3 penial glands with accessory flagella. Reproduction oviparous, producing pelagic egg capsules, plano-convex in shape. The capsule measured about 350m in maximum diameter and 170m in height, the egg about 84 in diameter, there being usually only 1 egg per capsule (Kojima, 1957). The breeding season was found to be from Febru- ary to April at Asamushi, Japan. Measurements (mm)— length width no. whorls locality 22.8 16.0 4 Hakodate, Japan 20.1 13.7 4+ Kamoito, Teshiwo, Japan 19.3 15,2 4 Muroran, Japan 17.8 13.3 4 Muroran, Japan 16.2 12.0 4 Awaji, Japan 14.4 11.4 4 Muroran, Japan 12.6 9.2 5 Shirahama, Japan 11.2 8.5 3 Side Saddle Island, China 10.8 8.5 4 Cheetfoo, China 9.3 6.8 4 Pei-tai-ho, China Synonymy— 1844 Turbo (Litorina) breviculus Philippi, Zeitschrift fur Malakolozoologie, vol. 1, p. 166 (China, at mouth of Yang-tse-Kiang (Largilliert!); 1847, Abbildungen und Beschreibungen Conchylien, vol. 2, p. 161, Litorina, pl. 3, fig 10. (lectotype figure). 1857 Littorina balteata Reeve, Conchologia Iconica, vol. 10, Littorina, pl. 14, fig 71, a,b (China; lectotype BM(NH) 1968316: 11 x 7.6 mm.). 1861 Litorina mandshurica Schrenck, Bulletin de L’Académie Impériale des Sciences de St. Pétersbourg, vol. 4, no. 7, p. 409 (Sinus des Castries [Zaliv Chikhacheva, Gulf of Tartary, USSR]) type in Academy of Sciences Leningrad? 1867, Reisen und Forschungen im Amur-Lande, vol. 2, p. 333, pl. 14, figs 14-20. 1862 Littorina souverbiana Crosse, Journal de Conchyliologie, vol. 10, p. 53, pl. 1, figs 6-7 (in Sinu Talienwhanensi [North China]; holotype MHNP: 13 x 10 mm), 1867 Litorina mandshurica carinata Schrenck, Reisen und Forschungen im Amur-Lande, vol. 2, p. 338 (no locality given). 1867 Litorina mandshurica laevigata Schrenck, ibid. 1887 Littorina manschurica ‘Schrenck’ Tryon, Manual of Conchology, vol. 9, pp. 241, 300 [error for L. mand- shurica Schrenck]. 1928 Littorina heterospiralis Grabau and King, Shells of Peitaiho, Peking Society of Natural History, Hand- book no. 2, second ed., p. 230, pl. 10, fig. 104 (Peitaiho [China]); types in Peking? Types—The type specimen of Turbo (Litorina) breviculus Philippi may be in the Berlin Muse- um, although it has been impossible for me to determine this with certainty. The lectotype fig- ure appears in Abbildungen und Beschreibungen Conchylien, pl. 3, fig 10 (see our pl. 329 figs. 5, 6). The lectotype of Littorina balteata Reeve is in the British Museum (NH)1968316. Types of L. mandshurica Schrenck and of its subspecies car- inata and laevigata, may be in the Academy of Sciences, Leningrad. The holotype of L. souver- biana Cross is in the Paris Museum (Journal de Conchyliologie Collection). The type of L. heter- ospiralis Grabau and King, may be in Peking. Records—KOREA: Wonsan Beach, Wonsan (MCZ); 2 mi. W. of Masan (USNM); Sorai (Sorai-men) (ANSP). CHINA: Port Arthur (Lu-shuh), Manchuria (ANSP, MCZ); Pei-tai-ho, N. Chihli coast; Cheefoo (Yen T’ai); Chiaochou (Kiautschou), nr. Tsingtau (Isington), Yellow Sea; Side Saddle Island (Lu- Hua Shan), Che-Kiang Province; Foochow (Fu-chou) (all USNM); Wuchiu Hsu (ZMC). JAPAN: HONSHU: Kominato; Yenosima (Islands) (USNM, MCZ); Isu, Ise (Isu-shima?) (MCZ):; Tokyo Bay (Yedo Bay) (ANSP, MCZ); Tsukishima, Tokyo, Tokyo Bay (USNM, MCZ); Shirahama, Wakayama Prefecturate; Yokohama (both ANSP, USNM); Misaki, Sagami Bay, off Hondo (USNM); Sagami Bay, Zushi Beach (MCZ); Shinagawa Bay, Tokyo; Toshima, Tanabe-wan, Seto Marine Biology Lab., Shirahama, Wakayama-ken (both USNM); In- land Sea (Seto-Naikai) (MCZ, ANSP); Awaji-shima (Island) (USNM, MCZ); Wakanura; Takami (both USNM). KYUSHU: Nagasaki; Camp Mower, Sasebo (both USNM); Hirado, Hizen (ANSP, USNM): Shimonoseki; Saigo, Dogo Islands, Oki Group, Japan Sea, shore (both USNM). HOKKAIDO: Muro- ran (USNM); Hakodate (USNM, MCZ); Zenibako (BPBM); Otaru; Kamoito, Teshio (both USNM). i aoe ae \4o0° 160° f i “ 4 PHILIBPINE “Guar of 7, a 3 Enwefok. > ce J) LU? | Oth Bhs Wg ee ‘ . . . > ws ee : CAROLINE ae te Plate 331. Geographical distribution of Littorina brevicula (Philippi.) sl [05-286] November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 435 Subgenus Littoraria Griffith and Pidgeon, 1834 Type: Littorina zebra (Donovan, 1825) Members of this subgenus are inhabitants of tropical and subtropical portions of the Indo- Pacific, East Pacific and West Atlantic, but repre- sentatives appear to be absent from the East Atlantic. Although the type-species of Littoraria is from tropical west America, the majority of species live in the Indo-Pacific with one species in the Caribbean area. This may reflect what appears to be a trend toward a higher rate of speciation in the Indo-Pacific. In this group the species generally are of medium size and are moderately robust; they are usually marked with axial or zigzag color lines, or are of a distinct solid color. Apertural white lines, so common in certain species of Littorinas.s. and particularly of Austr- olittorina, Nodilittorina and Granulilittorina are usually absent or diffuse in Littoraria. All species probably produce a pelagic egg capsule, although observations on reproductive habits have been made on only a few. Possible evidence of a common lineage is given by the presence in most species of a basal flap on the penis; the flap contains no penial glands or “klebdrusén” as it does in other littorines. Synonymy— 1834 Littoraria Griffith, and Pidgeon, The Animal Kingdom, vol. 12, Mollusca and Radiata, Index, p. 598, pl. 1, fig 3. Type-species by monotypy: Littoraria pulchra ‘Gray’ (Sowerby, 1832) [= Littorina zebra (Donovan, 1825)]. Key To Principle Recent Species of Indo-Pacific Littoraria Leo ohell various ly Color patterned. sca. au io ad oo waves vd Soe ew hw hE a eS Ee Ld 2 Shell without color pattern, plain pinkish gray .............. coccinea p. 05-301 22 CC orumella wenerally: Dine to ViOlet -<..66 saad seGede evades Der ve Oe a4 woes ee 2M OMUeIIe WANTS ~ a soca ke ad ote ay Ace ew, bese Oe Od Owe OS ak: ace era hh gig d ews 4 3a. Color pattern of brown undulate fammules or spots 3b. Color pattern diffuse, whorls encircled with faint gray band 3c. Color pattern of rather straight, closely-spaced brown axial lines 4. Color pattern a “herring-bone”’ effect, shell, white apertural band present .. . re eee undulata p. 05-298 . mauritiana p. 05-304 .. kraussi p. 05-306 spire less than half the length of 586 ogee 3 eee ace Be praetermissa p. 05-307 4. Color pattern of closely-spaced brown punctations, spire more than half the length of shell, no white apertural band Ee ee ee pintado p. 05-297 [05-297] 436 Littoraria Littorina undulata Gray, 1839 (Pls. 325, 332, 333) Range—From Madagascar to western Polyne- sia; apparently rare in Hawaii; not reported from southeastern Polynesia nor from east Africa. Remarks—Due to its variability some forms of Littorina undulata are rather easily confused with several other Indo-Pacific species: L. scabra and L. coccinea, L. mauritiana and L. kraussi. From L. scabra it differs markedly in the anatomy of external reproductive structures, a large penile basal flap being present in undulata, whereas in scabra the entire base of the penis is thickened; scabra reproduces ovoviviparously while undul- ata produces a pelagic egg capsule; scabra usu- ally inhabits mangroves and other wooden struc- tures such as pilings, but undulata is usually Joseph Rosewater Littorinidae found on rocks; the shell of undulata is smaller (24 mm.) at maturity than scabra (over 40 mm.); although color patterns may appear superficially similar, upon close examination it may be noted that in scabra axial color markings are applied in a mosaic, the interruptions caused by the deeply incised spiral sculpture, while in undulata the markings appear continuous. The relationship is closer between undulata and the three other species, coccinea, kraussi and mauritiana. The features of all four are somewhat similar, and indeed they are considered members of the same subgenus, Littoraria. In the case of these species differences from undulata are more subtle and involve character of and presence or absence of color pattern, sculptural differences, relative proportions and geographical ranges (see species descriptions). x 12.6 mm.). Figs. 3,4. Lectotype of L. tenuis Philippi from Ticao Island, Philippines (BM(NH) 1968276; 14.7 x 8.4 mm.). Figs. 5,6,7,8. L. acuminata Gould from Mangsi Island [Philippines] (figs. 5,6: Holotype, USNM 5638; 8.9 x 4.8 mm.; figs. 7,8: paratype, USNM 612325, 6.8 x 3.7 mm.). Figs. 9-16. L. undulata from various localities: figs. 9,10. Pulau Stupai, Mentawai Ids., S.W. Sumatra (USNM 655218; 17.8 x 10.8 mm.). figs. 11,12. Pulau Siburu, N. of Sipora, Mentawai Ids., $.W. Sumatra (USNM 654710; 12.2 x 7.3 mm.); figs. 13,14. Bikini, Marshall Islands (USNM 585104; 20.05 x 11.8 mm.). figs. 15,16. Hien- ghene, New Caledonia (USNM 666080; 17.3 x 10.05 mm.). Fig. 17. Lectotype figure of L. columna ‘Jonas’ Philippi, from Abbildungen und Beschreibungen Conchylien, vol. 3, pl. 6, fig. 15 (about 18 x 11 mm.). [05-298] November 30, 1970 The western Atlantic Littorina nebulosa (Lamarck) is the analogue of the Pacific L. undu- lata. Habitat—Lives intertidally on rocky shores and commonly clusters in rock crevices during day- light hours, becoming active at night and crawl- ing about on the rocks. Often associated with L. coccinea (Gmelin) where their ranges overlap. Description—Shell reaching 23.9 mm. (about .9 inch) in length, turbinate in shape, average obesi- ty about .58 (24 specimens range from .50 to .64) moderately thick in structure, imperforate, and sculptured with spirally incised lines, overall microscopic wavy spiral threads and fine, but rather regular oblique axial lines of growth. Ex- ternal color very variable: from a mottled yellow- ish gray to banded dark-brown, most often with continuous to interrupted undulating brown col- or markings; spire whorls often with white flam- mules on upper half, solid brown below; aperture light yellowish brown to dark yellowish brown sometimes with outside color markings showing through; columella usually grayish violet but sometimes lacking this color and being white, edged with brown. Base not flattened, separated from upper part of body whorl by a weak keel at periphery. Whorls 5-9, usually rather well round- ed. Spire less than half the length of shell, convex, produced at an angle of about 58°. Aperture broadly oval; outer lip thin in young individuals to moderately thick in adults; inner lip strongly produced, hardly concave, oblique, glazed with a grayish violet callus, pinched where it joins outer lip at base of columella. Suture impressed. Sculp- ture consisting of from 7-10 spiral striae on spire whorls, persisting on entire surface of body whorl where as many as 16 may be present above weak keel, and about the same number below; on body whorl somewhat weaker secondary striae may occur between primary ones especially at suture and near periphery. Entire surface covered with microscopic, closely-spaced wavy spiral threads not easily detectable in worn specimens. Axial sculpture consisting of rather regularly-spaced, oblique growth lines which upon crossing spiral sculpture produce a weak reticulation. Opercu- lum corneus, paucispiral. Periostracum not evi- dent in specimens examined. Nuclear whorls partially decollate in all specimens examined; remaining portions smooth and glassy; first 2-3 post-nuclear whorls light-brown and sculptured with spiral striae. Radula typically littorinid (2- 1-1-1-2). INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Littorina 437 Animal darkly pigmented on upper surface of tentacles, snout and foot. Verge with a large flap at its base, long and slender, but constricted about half its length and with a delicate translucent distal extremity. Seminal duct deeply folded. Re- production oviparous; egg capsules observed to contain single ovum; capsule simple, convexo- convex, with flotation “‘skirt.”” Capsule diameter ranging from .23-.25 mm., ovum diameter .09-.11 mm. Development undoubtedly pelagic; well-de- veloped veligers remaining in capsules on fifth day (Eniwetok-February) after shedding of cap- sule; pelagic stage probably long lived. Measurements (mm)— length width no. locality whorls 93.9 11.9 8+ Pulau Nias, $.W. Sumatra 21.8 12.0 6+ Guadalcanal, Solomon Islands 19.8 10.6 6+ Guadalcanal, Solomon Islands 19.7 11.6 6+ S.W. tip Sanding Id., Mentawai Ids. 18.0 10.3 6+ Guadalcanal, Solomon Islands 17.4 TT 5+ Bikini Atoll, Marshall Islands L5.7 9.9 6+ Bikini Atoll, Marshall Islands 14.5 7.8 7+ Pulau Nias, $.W. Sumatra 11.3 6.5 8 Pulau Nias, S.W. Sumatra 10.9 6.1 8 Pulau Nias, S.W. Sumatra Synonymy— 1839 Littorina undulata Gray, The Zoology of Captain Bee- chey’s Voyage—in His Majesty’s Ship Blossom, Mol- lusks, p. 140 (no locality given; not figured); type locality here designated: Okinawa, Ryukyu Islands; Lectotype BM(NH) 1968362, British Museum (N.H.). Plate 333. Littorina (Littoraria) undulata (Gray) Fig. A. Radula (upper scale is 0.1 mm.). Fig. B. Penis (lower scale is 2 mm.). Fig. C. Egg capsule from above. Fig. D. Egg capsule from side to show flotation “skirt” (capsule diameter about 0.23 mm.); all from Eniwetok, Marshall Islands. 438 Littoraria 1846 Littorina tenuis Philippi, Proceedings of the Zoological Society of London (1845) p. 141 (Ticao Island, Phil- ippines); 1847 Abbildungen und Beschreibungen Conchylien, vol. 3, p. 18, pl 6, fig 8; lectotype and two paralectotypes, British Museum (N.H.) 1968276 1847 Litorina columna ‘Jonas’ Philippi, Abbildungen und Beschreibungen Conchylien, vol. 3, p. 14, Litorina pl. 6, fig. 15 (no locality given). 1849 Littorina acuminata Gould, Proceedings of the Boston Society of Natural History, vol. 3, p. 84 (Mangsi [Mangsee] Id. [Balabac Strait, Philippines]); 1852, U.S. Exploring Expedition, Vol. 12, p. 200, pl. 14, fig 239 a, b, Holotype: USNM 5638. 1878 Litorina scabra concolor Weinkauff, Systematisches Conchylien-Cabinet, vol. 2, part 9, p. 37, pl. 4, fig. 11 [indicated in text]; in part (figs 8-10 are L. scabra) (Indo-Pacific). 1885 Littorina undulata contracta Nevill, Hand List of Mol- lusea in the Indian Museum, Calcutta, part 2, p. 143 (Andamans). 1885 Littorina undulata sulcatula Nevill, ibid.; p. 144 (Ascen- sion Island, Pacific [Ponape, Caroline Ids.]); refers to Reeve, 1857, Conchologia Iconica, vol. 10, Littorina, pl. 13, fig. 67a [occasional specimens approach this degree of rugosity]. 1885 Littorina conica subintermedia Nevill, ibid., p. 150 (Port Canning and False Point [Bengal, India]). Types—From the 3 syntypes of L. undulata in the British Museum (NH), catalogue number BM(NH) 1968362, the one most closely approxi- mating Gray’s measurements and description is here designated as lectotype (length 22.8, width 12.6 mm.; see our pl. 332 figs. 1, 2). Gray listed no type locality, but there are few places where the Blossom touched land within the range of this species (see Beechey, 1831; Rosewater, 1968). One such place, Okinawa, Ryukyu Islands, is here designated as the type locality. The speci- men of L. tenuis Philippi BM(NH) 1968276, figured in Abbildungen und Beschreibungen z0° 40° 60° 80° \oo° _ 1z0° Joseph Rosewater Littorinidae Conchylien pl. 6, fig 8, is here designated lecto- type (14.7 x 8.4 mm.; see our pl. 332 figs. 3, 4). The type specimen of L. columna Philippi may be in the Berlin Museum. The types of L. con- tracta, subintermedia and sulcatula Nevill should be in the Indian Museum, Calcutta. The holotype of L. acuminata Gould is in the U.S. National Museum (USNM 5638). The location of the type of L. concolor Weinkauff is unknown to me. Records—MADAGASCAR: Ambodifototra, Ile Ste. Marie; Ile aux Nattes, S. of Ile Ste. Marie; Faty, 13 mi. N. of Tulear (all MCZ); MAURITIUS: Pt. d’Espy N.E. of Poste de Flacq; E. of Souillac on Savanne River (both ANSP); SEYCHEL- LES: Anse a la Mouche, Mahe Island (YPM); MALDIVES: South half Kendikolu Island, Miladummadulu Atoll; btwn. Mafilefuri and Maro Islands, Fadiffolu Atoll; Dunidu Island, N. of Male, North Male Atoll; Ari Atoll (all USNM, ANSP); Hitadu Id., Haddumati Atoll (YPM); Wala Island, South Nilandu Atoll; N.W. tip of Gan, Addu Atoll (both ANSP). INDIA: Vengurla, N. of Goa; Goa; Khumpta (Kumta), N. Kanara (all USNM); Cochin, Malabar (MCZ, AMS); Cape Comorin; Remen Point, west side Pamban Pass, btwn. Gulf of Mannar and Palk Strait (both ANSP, USNM): Mandapam Camp (USNM); Madras (ANSP, MCZ). CEYLON: shore, Ft. Frederick, Trincomalee (ANSP, YPM); Weligama Bay; Pointe de Galle; Galle (all ANSP); Columbo (USNM, MCZ, NMW, SMF); Delft Island; Kankesanturai; W. of Kankesanturai; Merissa Village (all ANSP). MALAYSIA: Pulau Anyut, Malac- ca Strait, just S.W. of town of Malacca (USNM). THAILAND: Goh Huyong, Similan Islands; Goh Phi Phi (both USNM). FORMOSA: (ANSP). JAPAN: Higo (Hiigo Shima) (USNM); Hachijo Island, 275 mi. S. of Tokyo, Honshu (ANSP); Kago- shima (USNM): Osima, Osumi (USNM; MCZ). RYUKYU ISLANDS: Okinoerabu-shima (ANSP); 1 mi. N. of Shana Wan, Okinawa Island (USNM); Nago, Okinawa Island (BPBM, MCZ); Awa (ANSP). PHILIPPINES: Many locali- ties, including the following islands: Batanes; Babuyanes; Luzon; Burias; Mindoro; Panay; Cebu; Bohol; Mindanao; Basilan; Palawan (all USNM). SULU ARCHIPELAGO: Jolo Island; Simaluc Island, Tataan Islands, Tawi Tawi Group; Marongas Island, Jolo (all USNM). BORNEO: Sipitang; West Marudu Bay; Taganak (all USNM). CELEBES: Macassar (MCZ). MOLUCCAS: Karakelong Island, Talaud Islands (MCZ); Morotai, Halmahera Group, (USNM); Kasiroeta Is- e' UNDULATA | COCOE - KEELING ATOLLS Ow 20°
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Plate 334. Geographical distribution of Littorina (Littoraria) undulata (Gray).
[05-300]
November 30, 1970
land; Salangadeke Island, N. Loloda, W. of Halmahera Island
(both MCZ); Ceram (USNM); Hajara, Manipa Island (MCZ).
SUMATRA: Pulau Melila, S. of Udjung Batu, Banjak Islands
(USNM); Pulau We (MCZ, ZMA); Pulau Boenta, off Acheh
Head; Pulau Nias (both USNM); Padang (MCZ); Pulau Sibu-
ru, N. of Sipora; Veeckens Bay, South Pagi Island; Pulau
Stupai, north edge Sanding Island, Mentawai Islands; south-
west tip of Sanding Island, Mentawai Islands (all USNM);
Marong (ANSP). JAVA: Pelabuhan Ratu, Praeger (MCZ;
USNM); Welkomst Bay, Banten (USNM); Pemeungpeuk, nr.
Garut, S.E. coast of Java (MCZ). BALI (AMS). FLORES ID.
(ZMA). COCOS KEELING: North Keeling, near Pulo Selwa;
Cocos Lagoon; “Coral Pits,’ Klapa Tuju, West Island (all
USNM).
WESTERN AUSTRALIA: Broome; Ridell’s Beach, 2 mi. S.E.
of Gantheaume Point, Broome; Gantheaume Point, Broome
(all ANSP); S. end South Pasco Island, Barrow Island Group,
off Onslow (WAM, USNM). NEW SOUTH WALES: nr.
Wollongong (USNM); Port Stephens (AMS). QUEENS-
LAND: Heron Island, Capricorn Group (AMS); Cape Cleve-
land, nr. Townsville (USNM); Palm Island (AMS); Brook
Island (ANSP); Flying Fish Point; Russell River; Fitzroy
Island; nr. Cairns; Lizard Island (all AMS); Green Island, nr.
Cairns (MCZ); Cooktown; Murray Id., Torres Strait (both
AMS). NEW GUINEA: Hollandia; Aitape; outlet of Kalveng
River, Gusika, 13 mi. N. of Finschhafen (all MCZ); Huon
Gulf (USNM); Collingwood Bay (AMS); Yule Island (USNM;
BPBM:; ANSP; MCZ): Island 1/2 mi. S.S.W. OF Sowek,
Soepiori, Schouten Islands (MCZ); Biak; N.E. end of Noekori
Island; E. Noesi Isle, Mios Woendi Atoll, Padaido Islands;
Rainbawi Point, Japen Island; S.E. entrance to Wooi Bay,
Japen Island; Montoewoeri Point, Koeroedoe Island; Abroeki
Island, Maransabodi Island, Aoeri Islands, Geelvink Bay (all
ANSP). ADMIRALTY IDS.: Koruniat Island (ANSP). BIS-
MARKS: New Ireland (MCZ, USNM); Matupi Island, Rabaul,
New Britain (AMS). SOLOMONS: Woodlark Island (USNM);
Harapa, Shortland Island (MCZ); nr. mouth of Lunga River,
N. shore Guadalcanal (MCZ, ANSP); Point Purvis, Florida
Island; Ugi Island (both USNM); Makira Harbor, San Cris-
tobal (AMS). SANTA CRUZ IDS.: Tinakula (AMS). NEW
HEBRIDES: Pakea, Banks Group (AMS). LOYALTIES: Li-
fou (USNM; AMS); Maré (MCZ). NEW CALEDONIA: N. of
Hienghene (AMS); Koe Reef, 2 mi. $.S.E. Touho (ANSP); Isle
of Pines (MCZ); Ilot Maitre, about 4 mi. S.W. of Noumea;
(ANSP): Ilot Amédie (AMS); La Roche, Bourail; Gatope
Island, Voh (both ANSP). LORD HOWE ISLAND: Norfolk
Island (both AMS). FIJI: Point N. of Namu, NNE Viti Levu
(MCZ); Irvines, nr. Malaqereqere, S.W. Viti Levu (USNM);
Lakemba (BPBM). MARIANAS: Saipan (ANSP;
USNM;:NMW); Tinian (MCZ); Tumon Bay, Guam (BPBM).
PALAUS: Koror, Malakal Dock; Eil Malk (both ANSP); An-
gaur (BPBM, MCZ). HELEN REEF: N. end Helen Islands
(ANSP). CAROLINE ISLANDS: Tomil Harbor Yap Island
(USNM); Ulithi Atoll (ANSP); Elato Atoll (USNM); Dublon,
Truk; Ponape (both BPBM); Kusaie (AMS, BPBM, MCZ,
USNM); Kapingamarangi (USNM). MARSHALL ISLANDS:
Eniwetok; Bikini; Kwajelein; Arno; Jaluit; Ebon Atoll (ALL
USNM).
GILBERTS: Abaiang (MCZ); Maiana; Tabiteuea (both
BPBM); HAWAIIAN ISLANDS: Kailua, Oahu (BPBM); Ka-
milo, Kau; Keaukaha, Hilo, both Hawaii (both USNM). LINE
Islands; Palmyra (USNM). JOHNSTON ISLAND (USNM,
BPBM). ELLICE ISLANDS: Nui; Vaitupu; Nukualailai (all
USNM). WALLIS ISLANDS: Nukuhifala; btwn. Luanna and
Fungalei Islands (both USNM). HOORN ISLANDS: Futuna
(USNM). PHOENIX ISLANDS: Canton Island (USNM). SA-
MOA: Salelesi Village, Upolu Island (ANSP); Pago Pago,
Tutuila (BPBM; USNM). TONGA: Niuafoou Island; Niutoua,
Tongatapu; Laulea Reef, nr. Fatuma, Tongatapu (all USNM);
Velitoa; Tongatapu; Nomuka, Ha’apai (both BPBM).
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 439
Littorina coccinea (Gmelin, 1791)
(Pls. 325, 335, 336)
Range— Cocos-Keeling Islands to Eastern
Polynesia, including Hawaii.
Remarks—The name coccinea is derived from
the Latin ““coccineus” meaning “red like a berry”
and seems to be a misnomer in the case of adult
specimens which are largely whitish. There is,
however, always a blush of brownish orange
coloration which shows through and the aperture
is intensely colored. Young individuals show
more of the color which perhaps prompted Gme-
lin to adopt Martyn’s appellation and, in addition,
Martyn’s plate shows an exaggeratedly red-
colored invididual. Although evidencing no out-
standing characteristics the species is easily rec-
ognizable and in the center of its range, Microne-
sia, probably is the most common littorinid pres-
ent (see Demond, 1957).
Habitat— Lives intertidally on rocky shores
and commonly clusters in rock crevices during
daylight hours, becoming active at night and
crawling about on the rocks. Often associated
with L. undulata Gray where their ranges over-
lap.
Description—Shell reaching 25.8 mm. (slightly
over 1 inch) in length, elongate- to conic-
Plate 335. Littorina (Littoraria) coccinea (Gmelin).
Figs. 1,2. Specimen from Aitutaki, Cook Islands, showing
diffuse color-banding (USNM 684817; 14.3 « 8.1 mm.).
Figs. 3,6. Lectotype figure, from T. Martyn, Universal Con-
chologist, vol. 2, pl. 68 [external figs].
Figs. 4,5. Specimen from Vitilevu, Fiji, nearly white with
little evidence of external color or banding (USNM
531795; 22.7 x 14.0 mm.).
[05-301]
440 Littoraria
turbinate in shape, average obesity about .60, 21
specimens ranging from .56-.65, females slightly
more obese than males), older individuals rela-
tively thick in structure, imperforate, and sculp-
tured with spirally incised lines, overall micro-
scopic, wavy, spiral threads and irregular, ob-
lique lines of growth. External color white, lack-
ing patterned color markings except for occasion-
al circular white blotches on early whorls; with
brownish orange to strong brown within the
aperture and showing through on lower portions
of early whorls, on center of body whorl and on
the base of shell. Base only slightly flattened,
separated from upper part of body whorl by a
weak to moderate keel at the periphery. Whorls
6-9, moderately rounded. Spire less than half the
length of shell, convex, produced at an angle of
about 60°. Aperture roundly oval; outer lip thin in
young individuals to moderately thick in adults;
inner lip concave, curves obliquely posteriorly,
glazed with the brownish orange callus, pinched
where it joins outer lip at base of columella.
Suture distinctly impressed.
Sculpture consisting of from 7-10 spiral striae
over entire surface of early spire whorls, but often
becoming indistinct on penultimate and body
whorls where the first 1-5 striae only may be
visible; first subsutural stria the strongest, pro-
ducing a shouldered effect. Entire surface of shell
covered with microscopic, closely-spaced wavy
spiral threads not easily detectable in worn speci-
0.4mm
t =)
Plate 336. Littorina (Littoraria) coccinea (Gmelin), from
Eniwetok, Marshall Islands.
Fig. A. Radula (upper scale = 0.1 mm.)
Fig. B. Penis (lower scale is 5 mm.)
Fig. C. Two views of egg capsule (diameter of capsule
about 0.24 mm.; diameter of egg about 0.09 mm.).
Joseph Rosewater
Littorinidae
mens. Axial sculpture consisting of occasionally
coarse but usually fine oblique lines of growth.
Operculum corneus, paucispiral. Periostracum
not evident in specimens examined. Nuclear
whorls partially decollate in all specimens ex-
amined; remaining portions smooth and glassy;
first 2-3 post-nuclear whorls brownish orange and
sculptured with spiral striae. Radula typically
littorinid (2-1-1-1-2).
Animal darkly pigmented on upper surfaces of
tentacles, snout and foot. Verge long and slender,
with a swollen enlargement at its base. Reproduc-
tion oviparous; pelagic egg capsules observed to
contain single ovum; capsule simple, convexo-
convex, with flotation “skirt.” Capsule diameter
0.24; ovum diameter .09 mm. Development un-
doubtedly pelagic; veliger stage reached in 2-3
days (Eniwetok-February) after shedding of cap-
sules; some larvae swimming free.at that time.
Measurements (mm)—
length width no. whorls locality
25.8 15.0 6+ Keaukaha, Hilo, Hawaii
23.0 14.3 5+ Bikini Atoll, Marshall Ids.
20.4 12.0 9 Futuna, Hoorn Ids.
18.1 10.5 8+ Makatea, Tuamotu Ids.
15.3 9.2 8+ Makatea, Tuamotu Ids.
14.3 9.3 9 Makatea, Tuamotu Ids.
12.7 74 7 Huahine, Society Ids.
11.2 6.3 8+ Moorea, Society Ids.
9.8 5.8 7 Moorea, Society Ids.
7.2 4.2 6+ Moorea, Society Ids.
Synonymy—
[1788 Limax coccinea Martyn, Universal Conchologist, Vol. 2,
plate 68; rejected work, non-binomial. ]
1791 Helix coccinea Gmelin, Systema Naturae, edition 13, vol.
1, part 6, page 3651 (New Zealand [Tahiti, Society
Islands, here corrected]); refers to Martyn, Univ.
Conch. 2, t. 68, f. ext [restricted to external figures;
the figure on left (apertural view) here designated as
lectotype].
1832 Littorina obesa Sowerby, Genera of Recent and Fossil
Shells, vol. 32, no. 37, fig, 6 [fig 6 here designated as
lectotype] (South Sea Islands)
1839 Littorina limax Gray, the Zoology of Captain Beechey’s
Voyage in His Majesty’s Ship Blossom p. 139 (Society
Islands); refers to Martyn, Univ. Conch. [vol. 2, pl.
68, left external figure here designated as lectotype].
Types—Gmelin based the name Helix coccinea
on the figures in Martyn’s Universal Conchologist
(vol. 2, pl. 68, external figures). The location of
Martyn’s collection is not known and therefore
we must resort to his figures to distinguish this
fortunately distinctive species. The left hand fig-
ure, apertural view, is here designated as lecto-
type of Gmelin’s Helix coccinea. The type locali-
ty, given by Gmelin as New Zealand is here
corrected to Tahiti, Society Islands, a likely place
[05-302]
November 30,-1970
for Martyn’s figured specimen to have been col-
lected.
The location of type specimens of L. obesa
Sowerby and of L. limax Gray are also unknown;
they were not found in the collection of the
British Museum (N.H.) during a recent visit there
(1968). For L. obesa, Sowerby’s figure 6 is here
designated as lectotype; for L. limax, Martyn’s
left external ne pl. 68 of the Universal Con-
chologist, vol. 2, is here designated as lectotype.
Records—FORMOSA (ANSP). RYUKYU ISLANDS: Kikai-
ga-shima; Okinawa (both USNM). PHILIPPINES: Sabtan
Island; Santo Domingo, Batan Island, both Batanes Group;
Cujangan Island; Candaraman Island, Balabac (all USNM).
INDONESIA: Karakelong Island, Moluccas (MCZ). CO-
COS-KEELING ISLANDS: North Keeling (USNM). AUS-
TRALIA: Murray Island, Torres Strait; Lizard Island, (both
AMS); Green Island, nr. Cairns (MCZ); Herald Cays, off
Cairns; West Cay, Diamond Islets; Russell Island; Flying
Fish Point, all Queensland (all AMS), NEW GUINEA: 5 mi.
N.W. of Rani Island, Biak Island, Schouten Islands, (ANSP);
Hollandia (MCZ). BISMARCKS: Malie Island, Lihir Islands,
New Ireland (AMS). SOLOMONS: Stirling Isle, Treasury
Islands (USNM); Simbo Island (AMS, ANSP); Ugi Island
(USNM); Mortlock Ids. (Del. Mus. Nat. Hist.). SANTA CRUZ
ISLANDS: Vanikoro (AMS). NEW HEBRIDES: Espiritu
Santo Island (MCZ); Vate Island (Efate Island) (AMS); Tana
Island (USNM). LOYALTIES: Uvea Island (USNM); Lifou
Island (AMS). NEW CALEDONIA: N. of Touho (ANSP,
USNM); Ilot Maitre $.W. of Noumea (ANSP). LORD HOWE
ISLAND (AMS): FIJI: Cevu Island, Nandronga; Irvines, Nr.
Malaqeregere, both Viti Levu; Fulanga, Lau Group (all
USNM). MARIANAS: Saipan Island (ANSP, MCZ); Tinian
(USNM, MCZ); Tumon Bay, Guam (BPBM); Apra Bay, Guam
(USNM). PALAUS: S.E. Eil Malk; S.E. Auluptagel Island,
Malakal Harbor, Koror (both ANSP). CAROLINES: Yap
(BPBM); Mog Mog Ulithi (BPBM, eee: Ponape (MCZ);
Kusaie, “Coral Islet’ Lele (BPBM); Kapingamarangi (USNM,
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 441
BPBM). MARSHALL ISLANDS (many localities on the fol-
lowing atolls): Eniwetok; Bikini; Rongelap; Rongerik; Ujae;
Pokak; Bikar; Taka; Uterik; Wotho; Likiep; Ailuk; Arno; ie
Kwajalein; Ailinglapalap; Jaluit Atoll (all USNM). GIL-
BERTS: Abaiang Island (MCZ); Abemama Island (USNM):
Tabiteuea (BPBM). HAWAIIAN ISLANDS: Kaaawa,
USNM); Koko Head, western side, both Oahu (BPBM): Hilo,
(MCZ, USNM); Kapoho, (BPBM); Pohoiki, all Hawaii
USNM).
LINE ISLANDS: Kingman Reef (BPBM); Palmyra Island
(ANSP, USNM, AMS, MCZ, BPBM); Washington Island
BPBM); Fanning Island (BPBM, AMS); Flint Island (USNM,
ANSP); JOHNSTON ISLAND (USNM, MCZ, BPBM). EL-
LICE ISLANDS: Nanumea; Nui (both USNM); Funafuti
AMS, MCZ, USNM); Nukulailai (USNM); Rotuma (BMNH).
WALLIS ISLAND: Nukuhifala (USNM). HOORN _IS-
LANDS: Futuna (USNM). HOWLAND ISLAND (BPBM).
BAKER ISLAND (BPBM). PHOENIX ISLANDS: Canton
Island (USNM). SAMOA: Apia (USNM, AMS); Saluafata
reefs, both Upolu (ANSP); Pago Pago Harbor, (BPBM,
USNM); Aunuu Island, both Tutuila (MCZ); Ta’u Island
(BPBM, MCZ); Fakaofo Island (BPBM), Tokelau Ids. TON-
GA: Niuafo’ou (USNM; BPBM); Lifuka, Ha’apai Group;
Velitoa, (both BPBM); Popua Reef, Tongatapu (USNM).
COOK ISLANDS: North tip Akitua, Aitutaki (USNM); Taun-
ganui, Mauke (BPBM); Rarotonga (AMS); Avatiu Harbor to
Motu Tou, Rarotonga; Mangaia (both USNM, MCZ). AUS-
TRAL ISLANDS: Raeivavae, (USNM). SOCIETY ISLANDS:
Fanui Bay; S. of Farepiti Point; Vaitape V illage all Bora Bora;
around point from “Hipu”’; Bay Vaiore both Tahaa; Uturoa;
Il6t Tipaemanu S.E. of Utoroa; Tevaitoa, all Raiatea; Point
Teffaao, around reef, Huahine; Moto Fareone; Opunohu Bay
(all USNM); Pareau Point, District of Teavaro (ANSP), all
Moorea; Tiarei; btwn. Hamuta River and Fautaua River,
Pirae; Patutoa (all USNM); Atiue, Punaauia (ANSP); 43 kin.
N. of Pariauta, Hitiaa, (USNM); btwn. mouths of Maruapo
and Hapaa Rivers, District of Punaauia (ANSP); 1 km. S. of
Faaone River, Faaone; Motu Fenuaina, Tautira (both USNM)
all Tahiti. TUAMOTUS: Tikehau; Vahitahi; Reao; Tureia;
Makemo (all USNM); Takaroa; Niau; Toao; Amanu (all
ANSP): Makatea; Fakarava; Raroia (all ANSP; USNM). GAM-
BIER ISLANDS: Mangareva (MCZ; USNM: AMS). Oeno
Island; Pitcairn Island (both USNM). HENDERSON IS-
LAND (Smith, 1913).
60° 80° too® 120° 1408 160° 180° 160 - 140 120 100 90
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Plate 337. Geographical distribution of Littorina (Littoraria)coccinea (Gmelin),
[05-303]
442 Littoraria
Joseph Rosewater
Littorinidae
Littorina mauritiana (Lamarck, 1822)
(Pls. 325, 338, 339)
Range— South Africa and Southwestern Indian
Ocean Islands.
Remarks—Littorina mauritiana is restricted in
its distribution to the southwestern Indian Ocean.
It has been confused quite often with similar
appearing species from South Africa, Australia
and New Zealand but is unquestionably distinct
when its peculiarly smooth gray banded shell and
diffuse markings are recognized. The species
most commonly confused with L. mauritiana is
L. unifasciata and its subspecies in Australia,
New Zealand and Juan Fernandez, but it is easily
distinguished on the basis of shell and animal
characters. This species is closely related to Litto-
rina coccinea of the Western Pacific and also L.
kraussi of the Indian Ocean.
Habitat—Intertidal on rocks.
Description—Shell reaching 24 mm. (about 0.9
inch) in length, turbinate in shape; average obesi-
ty about .59 (13 specimens range from .55 to .63),
older individuals relatively thick in structure,
imperforate; sculptured spirally with overall mi-
croscopic wavy threads and axially with occa-
sional lines of growth. External color pattern
rather constant, consisting of a spiral band of gray
on middle of body whorl and on anterior third of
spire whorls; area anterior and posterior to band
with indistinct wavy axial color lines; anterior
color lines ending at another rather indistinct
gray spiral band on the base of body whorl with
only occasional subsequent, sometimes obliquely
spiral, color markings; overall background color a
yellowish gray. Aperture usually dark brown,
often with outside color pattern showing through;
with an internal basal white spiral band and
occasionally a small one posteriorly inside outer
lip. Whorls 6-8 but spire often eroded. Spire less
than half the length of shell, convex, produced at
an angle of about 57° (but varying to as low as
48°). Aperture oval; outer lip thin in young speci-
mens to moderately thick in adults; inner lip
concave, curving obliquely posteriorly, glazed
with a brown to violet callus, pinched where it
joins outer lip at base of columella. Suture im-
pressed. Sculpture consisting of microscopic,
closely spaced, wavy spiral threads not easily
detectable in worn specimens. Axial sculpture
consisting of irregular lines of growth which may
Plate 338. Littorina (Littoraria) mauritiana (Lamarck)
Figs. 1,2. Lectotype of P. mauritiana from “‘Tle-de’France,”
(MHNG 1096/91-2, 22.8 x 13.2 mm.).
Figs. 3,4. Paralectotype (MHNG 1096/91-5, 12.5 x 7.4
mm.).
Figs. 5,6. Lectotype of Littorina laevis Philippi, (BM(NH)
1968221, 18.4 x 11.4 mm.).
Figs. 7,8. Holotype of Tricolia vitrea Deshayes from Bour-
bon [Reunion] (MHNP, 6.1 x 4.2 mm.).
Figs. 9,10. L. mauritiana from Pointe Fayette, E. Mauriti-
us, (USNM 637330 [ex ANSP 273712] 19.8 x 11.1 mm.).
Figs. 11,12. L. mauritiana from E. of Souillac, Mauritius,
(USNM 637331 [ex. ANSP 274078] 15.8 x 9.4 mm.).
(Figs. 1,2,7,8. R. Robertson photos; other photos by the
author)
have associated dark pigmentation. Operculum
corneus, paucispiral. Periostracum not evident in
specimens examined. Nuclear whorls partially
decollate in all specimens examined; remaining
portions smooth and unsculptured. Radula typi-
cally littorinid.
Animal darkly pigmented on upper surfaces of
tentacles, snout and foot. Verge moderately long
and slender, with a terminally swollen branch
near its base, similar to L. kraussi. Reproduction
unknown; [probably oviparous and involving pe-
lagic capsules].
[05-304 ]
November 30, 1970
Measurements (mm.)—
length width no. whorls locality
24.0 13.3 7+ Mauritius
PA P22 6+ Mauritius
19.9 EL 8+ Mauritius
19.6 12.4 5+ Madagascar
18.9 10.8 5+ Mauritius
18.1 11.4 7+ Reunion
17.5 il 6+ Mauritius
17.4 9.8 6+ Mauritius
14.1 8.7 5+ Mauritius
11.4 7.0 6+ Mauritius
Synonymy—
1822 Phasianella mauritiana Lamarck, Histoire Naturelle des
Animaux sans Vertebres, vol. 7, p. 54 (Ile-de-France
[Mauritius]); Lectotype in Museum d Histoire Na-
1838 Phasianella mauriciana ‘Lamarck’ Potiez and Michaud,
Galerie des Mollusques, Muséum de Douai, vol. 1,
p. 311 (error for P. mauritiana Lamarck, 1822).
1846 Littorina laevis Philippi, Proceedings of the Zoological
Society of London (1845), part 13, p. 140 (locality
unknown [Mauritius here selected]); Lectotype in
British Museum (N.H.); BM(NH) 1968221, 18.4 x 11.4
mm. 1847, Abbildungen und Beschreibungen Con-
chylien, vol. 3, p. 10, Littorina, pl. 6, fig 6.
1847 Litorina mauritiana gracilior Philippi, Abbildungen und
Beschreibungen Conchylien, vol. 2, p. 165, Littorina,
pl. 3, fig 17b [right hand fig.] Probably is eastern
Pacific species best known as L. modesta paytensis
Philippi, 1847.
1863 Phasianella vitrea Deshayes, Catalogue des Mollusques
de L’ Ile de la Réunion, (Bourbon) p. 76, pl. 8, fig 8
(Bourbon; Holotype in Muséum d Histoire Naturelle,
Paris, 6.1 x 4.2 mm.).
Types—There are 5 syntypes of Phasianella
mauritiana in the Lamarck Collection of the
Plate 339. Littorina (Littoraria) mauritiana (Lamarck) from
Pointe Fayette, E. Mauritius (ANSP 273712)
Fig. A. Radula (upper scale is 0.1 mm.)
Fig. B. Penis (lower scale is 5 mm.)
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 443
Muséum d Histoire Naturelle, Geneva, represent-
ing three different species of Indo-Pacific Lit-
torinidae: L. mauritiana, L. kraussi, and L. uni-
fasciata. The last cannot have occurred at Mauri-
tius, which Lamarck clearly stated as type locali-
ty, and L. kraussi does not fulfill both require-
ments of Lamarck’s description: shell “bluish
white” and columella “violet blue.” Syntypes 2
and 5 (MHNG 1096/91-2, 5) do fulfill these re-
quirements and are here designated as Lectotype
and paralectotype respectively of Phasianella
mauritiana Lamarck (see our pl. 338, figs. 1-4)
The specimen of L. laevis which Philippi fig-
ured in Abbildungen und Beschreibungen Con-
chylien vol. 3, pl. 6, fig 6, is in the British
Museum (NH) BM(NH) 1968221, and is here
selected as lectotype of that species (see our pl.
338, figs. 5,6). The holotype of P. vitrea Deshayes
is in the Muséum d'Histoire Naturelle, Paris (see
our pl. 338, figs. 7, 8). The type specimen of L.
gracilior Philippi may still be extant in the Berlin
Museum.
Records—SOUTH AFRICA: Port Edward, Natal (ANSP).
MADAGASCAR: Flacourt, Fort Dauphin (MCZ). SEYCHEL-
LES: Praslin Island (Melville, 1909). MAURITIUS: S. side
Tombeau Bay, W. Mauritius; E. Tamarin Bay, W. Mauritius;
Pointe Fayette, E. Mauritius; E. of Souillac on Savanne River
(all ANSP); Port Louis (MCZ). REUNION: Sainte Pierre
(AMS). CHAGOS ARCHIPELAGO (Melville, 1909).
Plate 340. Geographical distribution of Littorina (Littoraria)
mauritiana (Lamarck) in the southwestern Indian Ocean.
[05-305]
444 Littoraria
Littorina kraussi new name Rosewater, 1970
(Pls. 325, 341)
Range—East coast of Africa, the Indian Ocean
islands and south India to Cocos-Keeling Atolls.
Remarks—The familiar name Littorina glabra-
ta Philippi, 1846, is unfortunately a secondary
homonym of Paludina glabrata Pfeiffer, 1828
(=Littorina neritoides (Linné)), the Mediter-
ranean species and, therefore, must be renamed.
The species is renamed for F. Krauss from whom
Philippi “stole” several South African species by
publishing his manuscript names first. Littorina
kraussi is closely related to L. mauritiana, undu-
lata and coccinea.
Habitat—On rocks near high tide line.
Description—Shell reaching 21.8 mm. (about
0.9 inch) in length, turbinate in shape; average
obesity about .58 (15 specimens range from .55 to
.62); older individuals relatively thick in struc-
ture, imperforate, and sculptured with spirally
incised lines, overall microscopic wavy spiral
Plate 341. Littorina (Littoraria) kraussi Rosewater.
Figs. 1,2. Lectotype, L. glabrata [= L. kraussi] from Natal,
South Africa (BM(NH) 1968220, 18.3 x 10.8 mm.).
Figs. 3,4. Paralectotype, same BM number, 18.4 x 11.1
mm.
Figs. 5,6. Tiladummati Atoll, Maldive Islands (USNM
672385, 17.7 x 10.5 mm.).
Figs. 7,8. The same, a smaller specimen showing more
intense coloration (11.4 x 7.4 mm.).
Figs. 9,10. Zanzibar (USNM 89411, 14.7 x 8.9 mm.).
Figs. 11,12. Near Poste de Flacq, Mauritius (USNM
637326, 15.5 x 8.7 mm.).
Joseph Rosewater
Littorinidae
threads and fine rather regular oblique axial lines
of growth. External color pattern rather constant,
consisting of moderately regularly spaced, ob-
lique, zigzagging, or chevron-shaped brown
markings superimposed on a lighter background;
(pale to grayish-yellowish pink ISCC-NBS num-
bers 31-32) spire whorls above penultimate whorl
often dark anteriorly and light posteriorly, par-
ticularly noticeable in small individuals; aperture
light to darker brown, sometimes with exterior
color markings showing through; columella vari-
able in color; ranging from dark brown through
violet to nearly white with a brownish tinge. Base
not flattened, separated from upper part of body
whorl by a weak to moderate keel at periphery.
Whorls 6-7 or 8 but spire often eroded. Spire less
than half the length of shell, convex, produced at
an angle of about 55°. Aperture oval; outer lip thin
in young specimens to moderately thick in adults;
inner lip concave, curving obliquely posteriorly,
glazed with a brown, violet or nearly white callus,
pinched where it joins outer lip at base of colu-
mella. Suture impressed. Sculpture consisting of
from 7-10 spiral striae on spire whorls and per-
sisting onto body whorl where 10-11 may be
present above the keel and about the same num-
ber below; striae evanescent in some specimens,
worn away or weak at center of whorls. Entire
Zo* 40° : 60° 890°
i YE Se FS
ANDAME
1s
NICOB/
18.
EYLON
cocos
|
ee
Reunion
80°
Plate 342. Geographical distribution of Littorina (Littoraria)
kraussi Rosewater in the Indian Ocean.
[05-306 ]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 445
surface covered with microscopic, closely spaced,
wavy spiral threads not easily detectable in worn
specimens. Axial sculpture consisting of rather
regularly spaced lines of growth. Operculum cor-
neus, paucispiral. Periostracum not evident in
specimens examined. Nuclear whorls partially
decollate in all specimens examined; remaining
portions smooth and glassy; first 1-2 post nuclear
whorls light brown and usually sculptured with
spiral striae. Radula typically littorinid and simi-
lar to L. mauritiana.
Animal darkly pigmented on upper surfaces of
tentacles, snout and foot. Verge long and slender
with a terminally swollen branch near its base,
appearing very similar to L. mauritiana. Re-
production unknown; [probably oviparous and
involving pelagic capsules. ]
Measurements (mm.)—
length width no. whorls locality
21.8 11.9 6+ Mauritius
21.6 12.4 7+ Mauritius
19.5 11.6 T+ Mauritius
Meo 10.5 7+ Zanzibar
16.9 9.3 6+ Madagascar
14.4 8.6 7+ Mombasa, Kenya
13.1 Uso 6+ Madagascar
12.5 6.9 7+ Mombasa, Kenya
10.2 6.3 7+ Mombasa, Kenya
9.3 D0 7 Madagascar
Synonymy—
1846 Littorina glabrata Philippi, Proceedings of the Zoologi-
cal Society of London (1845) p. 140 (Payta Peru; and
Cape Natal [South Africa] here restricted to second
locality). 1848, Abbildungen und Beschreibungen
Conchylien, Vol. 3, p. 62. Litorina pl. 7, fig 5.
Lectotype in BM (NH) 1968220; [Non Paludina gla-
brata Pfeiffer, 1828=Littorina neritoides (Linnaeus,
1758)]. 1848, Krauss, Die Sudafrikanischen Mollus-
ken, p. 103.
Types—Although it was necessary to supply a
new name for the secondary homonym, Littorina
glabrata Philippi, 1846, the type specimens are
the same for both L. glabrata and L. kraussi. A
lectotype is here designated from among the four
syntypes of L. glabrata in the British Museum
(NH) (BM (NH) 1968220, length 18.3, width 10.8
mm.; see our pl. 341, figs. 1, 2).
Records—SOUTH AFRICA: Miller's Point, False Bay
(ZMA); Coffee Bay, 1 mi. S. Umtata River; East London, at
mouth Nahoon River; Second Beach, Port St. John; Port
Edward; Margate, 11 miles south of Port Shepstone; Isipingo
(all ANSP; NMW): Durban; mouth of Umhlali River (both
MCZ). MOZAMBIQUE: Ilha da Inhaca, Delagoa Bay
(ANSP). TANZANIA: Kendwa Island, 4 miles E.S.E. of
Dar-es-Salaam (MCZ); Bungi, Kiwani Bay; Changa; E. side
Puopo, Tumbat Island (all ANSP). KENYA: Mombasa Island
(ANSP; NMW). ADEN: (RNHL; BM(NH); MHNP). MADA-
GASCAR: Anakao, 20.5 miles S. of Tulear; Soalary, 16 miles
S. of Tulear; 1 mile N.W. of St. Augustin, 14 miles S.E. of
Tulear; bay S. side Nosy Iranja, 32 miles S.W. of Nossi Bay;
Nosy Tany Kely, 4 miles S. of Nosy Be; Pte. Fievre, S. Nosy
Be; S. of Anivorano, 12 miles N. of Ambodifototra, W. coast
Ile Ste. Marie; Ambodifototre, Ile Ste. Marie; S.W. shore Ie
aux Nattes S. of Ile Ste. Marie; Flacourt, Fort Dauphin (all
R.W. Foster; ANSP; MCZ). SEYCHELLES: St. Pierre Islet;
Mahé Island (both YPM); Praslin Id. (NMW). MAURITIUS:
S.E. side Tamarin Bay; S. side Tombeau Bay; !/4 mile S.W. of
Cape Malheureux; Pte. Lafayette; 1 mile N.E. of Poste de
Flacq; Pte. Vacoas, 3 miles S. of Mahebourg; near Savanne
River E. of Souillac (all ANSP). REUNION: (MCZ).MALDIVE
ISLANDS: Faro Islet reef, N.W. of Filadu Island, Tiladum-
mati Atoll; between Mafilefuri and Maro Islands, Fadiffolu
Atoll; Dunidu Island, North Male Atoll;
Wala Island, South Nilandu Atoll (all ANSP); Hitadu, Had-
dummati Atoll (YPM); Gan, Addu Atoll (YPM; ANSP); Ari
Atoll (ANSP). CHAGOS ARCHIPELAGO: Egmont Atoll
(Melville, 1909). INDIA: Goa (USNM). CEYLON: Point de
Galle (ANSP; RNHL). COCOS-KEELING ISLANDS: Cocos
lagoon, North Keeling (both USNM).
Littorina praetermissa May, 1909
(Pl. 325, 343, 344)
Range—Tasmania, Victoria and South Aus-
tralia.
Remarks—The “neglected” littorine is a dis-
tinct and easily recognizable species inhabiting
the southeastern Australian region. Considering
the narrow range it occupies it is not surprising
that it escaped attention until named by May. The
Plate 343. Littorina (Littoraria) praetermissa (May)
Figs. 1,2. Low Head, Tasmania; showing light-brown nu-
clear whorls and distinctive color pattern (USNM 671215:
11.9 x 7.7 mm.).
Fig. 3. Lectotype, original illustration from Papers and
Proceedings Royal Society of Tasmania for 1908, pl. 6,
fig. 3; Tasmania.
Figs. 4,5. Port Fairy, Victoria; large specimen showing
considerable wear and/or corrosion (USNM 637353: 16 x
11 mm.).
[05-307]
446 Littoraria
relationships of L. praetermissa appear to lie
more closely with the species of the tropical
Indo-Pacific than they do with the other Aus-
tralian species, such as L. unifasciata, although
the radula of praetermissa is similar in the nar-
rowness of the central tooth. It may be dist-
inguished at once from L. unifasciata by its
coarser spiral sculpture, color markings, less
deeply impressed umbilical crescent, and con-
siderably more rounded whorls. The animals also
differ in details of verge anatomy.
Habitat—F ound in clusters on rock surfaces at
and above the high water mark. (Macpherson and
Gabriel, 1962).
Description—Shell reaching 16.7 mm. (about .7
inch) in length, globose-turbinate in shape, aver-
age obesity about .66 (23 specimens range from
.61 to .71); moderately thick in structure, im-
perforate, developing a rather narrow crescent-
shaped area adjacent to columellar callus; sculp-
tured with shallow and relatively widely spaced
spiral striae and somewhat irregular axial lines of
growth; shell surface rather uneven and bumpy.
External ground color of shell grayish white, with
usually prominent brown zigzag lines overall.
Aperture medium brown with a white band at
junction of outer lip and columella; with white
markings around inner margin of outer lip. Base
hardly flattened, separated from upper part of
body whorl by a distinct to obsolete keel at
periphery. Whorls 4-5, rounded. Spire consider-
ably less that half the length of shell, convex,
produced at an angle of about 72°. Aperture oval
in outline; outer lip moderately thick, having its
origin high on body whorl above keel so that in
some specimens keel enters aperture as a simu-
lated parietal tooth; inner lip nearly straight.
Columella yellowish to tannish white, moder-
ately well excavated, with a low tooth-like swel-
ling near junction with outer lip; also usually
with a narrow crescent-shaped area on base ad-
jacent to columellar callus. Suture distinct. Sculp-
ture consisting of from 15-17 sometimes indis-
tinct spiral striae on body whorl and from 7-11 on
spire whorls; striae fairly distinct in young speci-
mens to indistinct in old invidivuals; shell sur-
face between striae (i.e. spiral cords) relatively
flat; peripheral keel on body whorl moderately
formed to obsolete. Shell surface overall rather
uneven and bumpy. Axial sculpture consisting of
irregular oblique lines of growth. Operculum
corneous, rather thin, paucispiral, nucleus well
delineated. Periostracum not evident in speci-
mens examined. Nuclear whorls at least partially
Joseph Rosewater
Littorinidae
decollate in all specimens examined; remaining
portions light brown in color, smooth, about 3 in
number. Radula littorinid (2-1-1-1-2) central tooth
moderately wide; lateral large and heavy, outer
marginal with 11 denticles.
Available rather poorly preserved animals dark-
ly pigmented on surface of tentacles snout and
foot. Verge only moderately short and thick; hav-
ing a simple, truncate basal flap. Nothing is
known concerning reproduction and develop-
ment of this species; probably oviparous, spawn-
ing pelagic capsule.
Measurements (mm.)—
length width no. whorls locality
16.7 11.9 4+ South Australia
[pA 9.5 5+ South Australia
14.7 9.5 4+ South Australia
14,1 9.6 4 Port Fairy, Victoria
13.8 8.5 4+ Port Fairy, Victoria
13.1 8.7 5 Low Head, Tasmania
12.0 7.8 5 Low Head, Tasmania
a7 7.6 4+ Queenscliff, Victoria
9.6 6.3 4+ Port Phillip Heads,
Tasmania
9.0 6.3 4 Queenscliff, Victoria
Synonymy—
1909 Littorina praetermissa May, Papers and Proceedings of
the Royal Society of Tasmania for 1908, p. 57, pl. 6,
fig 3 (Tasmania; type deposited in Tasmanian Muse-
um, Hobart: 15 x 11 mm; probable paratypes in
Australian Museum, Sydney and in Museum of Com-
parative Zoology).
Records—SOUTH AUSTRALIA: Robe (NMW); Boat-
swain’s Point, Nr. Robe (USNM, ANSP, MCZ); Encounter
Bay; Adelaide (both ANSP). VICTORIA: Shelly Bay, San
Remo (BM(NH)); Phillip Island (MCZ); Sandringham, Port
Phillip (ANSP); Port Phillip (AMS); Queenscliff (USNM);
Port Fairy (ANSP, USNM, AMS, NMW). TASMANIA: east
coast, Tasmania (AMS, paratype); Burnie; Wineglass Bay;
Cape Sorell; Opossum Bay; Sandy Cape (all AMS); Low Head
(USNM, MCZ); Eaglehawk Neck (AMS, MCZ).
Plate 344. Littorina (Littoraria) praetermissa (May)
Fig. A. Radula (upper scale is 0.1 mm.).
Fig. B. Penis (lower scale is 10 mm.); both from Port Fairy,
Victoria, Australia, USNM 637353.
[05-308 ]
November 30, 1970
s
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be HEBRIDES .- a
, Ol:
hs See Filis
|
|
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NEW
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00° 120° 140° 160° 160°
Plate 345. Geographical distribution of Littorina (Littoraria)
praetermissa (May) in southern Australia.
Littorina pintado pintado (Wood, 1828)
(Pls. 325, 346, 347)
Range—Southwestern Indian Ocean, from
South Africa to Mauritius; in the Western Pacific
from Japan through the North Pacific Islands to
Hawaii.
Remarks—Littorina pintado evidences a very
peculiar discontinuous distribution. It occurs ina
rather isolated outpost in the southwestern Indian
Ocean apparently skips the mainland of southeast
Asia and the East Indies and becomes established
again in the Western Pacific from whence it is
distributed more uniformly eastward to Hawaii.
In the eastern part of its range it is recorded only
from the North Pacific, but in the Indian Ocean it
occurs far south of the equator. This sort of
interrupted distribution is difficult to explain un-
less the concept is accepted of a once more evenly
distributed species that for unknown reasons has
disappeared from the center of its former range.
The possibility of long range rafting or even ship
transport cannot definitely be ruled out however.
The species is apparently quite hardy and there is
a note with two specimens in the USNM collec-
tion indicating that they were brought from Ha-
waii to Los Angeles, California, where they lived
for a year without seawater; an almost identical
observation was made by Edmondson (1946).
A darker subspecies, L. pintado schmitti occurs
at Clipperton Island. An apparent analogue of L.
pintado is L. pullata Carpenter of the tropical
eastern Pacific. Littorina tessellata Philippi of the
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 447
tropical western Atlantic, L. cingulifera Dunker,
and L. cincta Quoy and Gaimard of New Zealand
initially appear similar to pintado but upon closer
examination are quite distinct.
Habitat—Lives intertidally on rocky shores and
commonly clusters in rock crevices during day-
light hours, becoming active at night and crawl-
ing about on rocks. Observed to be associated
Plate 346. Figs. 1-12. Littorina (Littoraria) pintado pintado
(Wood); Figs. 13-16. L. (Littoraria) pintado schmitti (Bartsch
and Rehder).
Fig. 1. Lectotype of Turbo pintado Wood, from Sandwich
Isl.; a rather attenuate specimen (BM(NH) 1968368, 17
mm. length.)
Fig. 2. Lectotype of Littorina serialis Eydoux and Souleyet,
from iles Sandwich (MHNP, 18.6 x 11.8 mm.).
Figs. 3,4. Lectotype of Litorina ambigua Philippi from
Insulae Sandwich (BM(NH) 1968314, 15.9 x 9.7 mm.).
Figs. 5,6. Specimen from Hilo, Hawaii (USNM 339401,
15.5 x 9.7 mm.).
Figs. 7,8. Specimen from Bikini Atoll, Marshall Islands
(USNM 585105, 13.4 x 7.6 mm.).
Figs. 9,10. Specimen from Ani Jima, Bonin Islands (USNM
621892, 15.2 x 9.1 mm.).
Figs. 11,12. Specimen from East of Souillac, Mauritius
(USNM 637354, 12.2 x 7.3 mm.).
Figs. 13,14. Holotype of Littorina schmitti Bartsch and
Rehder, from Clipperton Island, eastern Pacific; note dark
coloration in this male specimen (USNM 472547, 12.2 x
7.6 mm.; this is specimen figured in original description).
Figs. 15,16. Paratype of L. schmitti Bartsch and Rehder,
from Clipperton Island, eastern Pacific, female specimen,
somewhat corroded, but showing dark coloration (USNM
472546, 14.7 x 8.8 mm.).
[05-309]
448 Littoraria
Joseph Rosewater
with L. coccinea and L. undulata at Eniwetok
and Bikini Atolls, Marshall Islands (personal ob-
servations; USNM records) and with L. picta in
Hawaii (Whipple, 1965; Struhsaker, 1966).
Description—Shell reaching 20.6 mm. (0.8
inch) in length, rather conic-turbinate in shape,
average obesity about .60 (18 specimens range
from .57-.63); older individuals moderately thick
in structure, imperforate, and sculptured with
spirally incised lines, overall microscopic wavy
spiral threads and oblique axial lines of growth.
External color yellowish to purplish gray with an
overall pattern of closely spaced reddish-brown
streaks or spots; the latter usually arranged in
spiral series and only seldom appear axially coor-
dinated; the center of body whorl and anterior
half of spire whorls often darker in color. Aper-
ture dark brown, the interstices of outer color
pattern showing through as light lines inside
outer lip. Base hardly flattened, separated from
upper part of body whorl by a rather weak keel at
the periphery. Whorls 5-9, moderately rounded.
Spire less than half the length of shell, convex,
produced at an angle of about 62°. Aperture oval;
outer lip thin in young individuals to moderately
thick in adults; inner lip weakly concave to nearly
straight, its inner margin a white crescent rimmed
with dark brown, relatively stout posteriorly but
pinched where it joins outer lip at base of colu-
mella. Suture moderately impressed; whorls often
slightly constricted below suture. Sculpture con-
sisting of about 10-11 spiral striae over surface of
spire whorls; from 10-14 striae on body whorl
above apertural line and 16-20 below on the base;
extra striae often inserted between main striae or
Plate 347. Littorina (Littoraria) pintado pintado (Wood)
Fig. A. Radula (upper scale is 0.1 mm.).
Fig. B. Penis; note lack of branches, flaps or glands (lower
scale is 2 mm.).
Littorinidae
in worn specimens sculpture may be partially
obscured. Character of shell surface between
striae variable, usually flat but may be raised
rounded cords. Entire surface of shell covered
with somewhat diffuse microscopic, closely
spaced wavy spiral threads not easily detectable
in worn specimens. Axial sculpture consisting of
rather irregular growth lines which are often
associated with dark axial pigmentation streaks.
Operculum corneus, paucispiral. Periostracum
not evident in specimen examined. Nuclear
whorls partially decollate in all specimens ex-
amined, remaining portions smooth and glassy;
first 2-3 post-nuclear whorls dark brown and
sculptured with spiral striae. Radula typically
littorinid (2-1-1-1-2).
Animal darkly pigmented on upper surface of
tentacles, snout and foot. Verge simple, lacking
lateral protuberances, grayish white. Reproduc-
tion oviparous; pelagic egg capsule plano-convex
in outline, measuring between .16 and .2 mm. in
diameter, usually containing a single ovum mea-
suring between .08 and .1 mm. (see Ostergaard,
1950; Whipple, 1965; for details of spawning and
development see Struhsaker, 1966).
Measurements (mm.)—
length width no. whorls locality
20.6 1233 5+ Midway Id.
19.3 11.0 5+ Midway Id.
19.1 11.0 7+ Chichi Jima, Bonin Ids.
17.0 10.2 5+ Osima Osumi
15.9 10.0 7+ Hawaii
14.7 9.2 5+ Osima Osumi
14.2 8.8 6+ Mauritius
13.0 CD 7+ Hawaii
12.3 74 6+ Mauritius
10.1 6.2 6+ Osima Osumi
Synonymy—
1828 Turbo pintado Wood, Supplement to the Index Testaceo-
logicus, p. 20, pl. 6, Turbo fig. 34 (Sandwich Isl.);
lectotype BM(NH) 1968368.
1839 Littorina tenebrata ‘Nuttall’ Jay, A Catalogue of the
Shells in the Collection of John C. Jay, 3rd ed; p. 73
(Sandwich Isles) [nomen nudum]; 1847 [as Litorina
tenebrata ‘Nuttall’] Philippi, Abbildungen und Be-
schreibungen Conchylien, vol. 2, p. 203 [in synonymy
of L. pintado Wood].
1848 Litorina ambigua ‘Nuttall’ Philippi, Abbildungen und
Beschreibungen Conchylien vol. 3, p. 62, Litorina,
pl. 7, fig 6 (Insulae Sandwich); Lectotype in B.M.
(NH) 1968314.
1852 Littorina serialis Eydoux and Souleyet, Voyage Autour
du Monde sur La Corvette La Bonite, Zoologie, vol.
2. pl. 31, figs 34-36, (Iles Sandwich); lectotype in
Museum d’Histoire Naturelle, Paris.
1882 Litorina pindata Philippi in Weinkauff, Systematischen
Conchylien-Cabinet, vol. 2, part 9, pp. 63, 70 [error
for L. pintado].
1882 Littorina ambiqua Reeve in Weinkauff, ibid., p. 70 [error
for L. ambigua].
[05-310]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 449
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Plate 348. Geographical distribution of Littorina (Littoraria)
pintado pintado (Wood) in the North Pacific Ocean and in the
southwestern Indian Ocean and of the subspecies, L. pintado
schmitti (Bartsch and Rehder), from Clipperton Island, east-
ern Pacific.
Types—From 17 syntypes of Turbo pintado
Wood from the Gray collection, the largest is here
designated lectotype (see our pl. 346 fig. 1; length
17 mm.; BM(NH) 1968368. The figured speci-
mens of L. tenebrata ‘Nuttall’ Philippi and L.
ambigua ‘Nuttall’ Philippi, are here designated
lectotypes of these species. The location of the
figured specimen of L. tenebrata is not known,
but that of L. ambigua is believed to be in the
British Museum (NH) (BM(NH) 1968314; 15.9 x
9.7 mm.; (see our pl. 346 figs. 3, 4). The lectotype
of L. serialis Eydoux and Souleyet, here designat-
ed, is in the Muséum d Histoire Naturelle, Paris
(see pl. 346 fig. 2; 18.6 x 11.8 mm.).
Records—SOUTH AFRICA: Margate, 11 mi. S. Port Shep-
stone, Natal, (ANSP). MOZAMBIQUE: Punta Abril, Delagoa
Bay (ANSP). MADAGASCAR: S. of Anivorano, 12 mi, N. of
Ambodifototra, W. coast He Ste. Marie; $.W.shore of Ile des
Nattes, S. of Ile Ste. Marie; Flacourt, Fort Dauphin (all MCZ).
MAURITIUS: Pointe Fayette; E.S.E. of Souillac (both
ANSP); Souillac S. coast, E. of Souillac (ANSP; USNM).
REUNION: (MCZ, NMW). JAPAN: Osumi Islands (USNM).
RYUKYU ISLANDS: Shuri; Odomari, both Okinawa (both
USNM). PHILIPPINES: Santo Domingo, Batan Island, Ba-
tanes Group (USNM). BONIN ISLANDS: (ANSP, MCZ,
NMW): Ani Jima; Chichi Jima (both USNM). MARIANAS:
Agrigan; W. Pagan (both USNM); Saipan (ANSP, MCZ,
USNM); Tinian (MCZ; BPBM; USNM). PALAUS: Angaur
(ANSP; MCZ; USNM; SMF). WAKE ISLAND: (USNM;
BPBM: ANSP). MARSHALL ISLANDS: Eniwetok; Bikini;
Rongelap Atolls (all USNM). HAWAIIAN ISLANDS: Mid-
way (USNM, MCZ, ANSP); Namokulaa, Eastern Islet, Mid-
way Atoll; Pearl and Hermes Reef, Grass Island; Lisianski
Island (all BPBM); Laysan (USNM, BPBM); Gardner Island;
La Perouse Pinnacle, French Frigate Shoals (both BPBM);
Necker Island (BPBM; USNM); Nihoa Island (BPBM); Kama-
lino Bay, Niihau Island (USNM); and many localities on
Kauai, Oahu, Molokai, Maui, Lanai, Kahoolawe and Hawaii
(see map; ANSP; BPBM; USNM, MCZ). JOHNSTON IS-
LAND: Sand Island (USNM, MCZ, BPBM).
Littorina pintado schmitti
(Bartsch and Rehder, 1939)
(Pl. 346 figs. 13-16)
Range—Known only from Clipperton Island.
Remarks—Hertlein) and Emerson (1953)
pointed out that a number of marine mollusks
which are distributed mainly in the Indo-Pacific
have been found on Clipperton Island, a locality
which also has strong eastern Pacific faunal affin-
ities. Littorina schmitti was described by Bartsch
and Rehder as being related to L. pintado. The
degree of relatedness is considered here to be
definitely of subspecific rank, the very distinct
differences between pintado and schmitti appa-
rently having been caused by geographic isola-
tion.
Habitat—Living intertidally on rocks.
[05-311]
450 Littoraria
Joseph Rosewater
Littorinidae
Description—Largest available specimen 18.5
mm. (about 0.7 inch) in length, average obesity
about .58 (20 specimens range from .54 to .64),
somewhat shorter than L. pintado, similarly
shaped and sculptured, but with darker colora-
tion, the individual color spots tending to be
larger and to coalesce, so that many specimens
appear to have a dark background with light spots
instead of the reverse as in L. pintado. Spiral
striae usually 7-10 on spire whorls and on body
whorl above apertural line, indistinct on the base;
shell often marked with white spiral lines, both
externally and within aperture; white lines often
coincide with striae. Inner lip nearly straight
when viewed from aperture; when viewed from
aspect of outer lip, columella is thickened and
pronouncedly convex at its midpoint.
Measurements (mm.)—
length width no. whorls locality
18.5 10.1 5 Paratype
18.4 9.9 4+ Paratype
lrg 10.5 4+ Paratype
15.7 9.0 5 Paratype
14.9 9.0 6 Paratype
13.9 8.2 5 Paratype
13.0 7.8 5 Paratype
12.6 7.4 7 Paratype
12.2 7.6 6+ Holotype
10.3 6.6 5 Paratype
Synonymy—
1939 Littorina schmitti Bartsch and Rehder, Smithsonian
Miscellaneous Collections, vol 98, no. 10 (Publica-
tion 3535), p. 9, pl. 2, fig 4 (Clipperton Island);
Holotype: USNM 472547; original measurements of
holotype (11.8 x 8.2 mm.) different from those ob-
tained here: 12.2 x 7.6 mm.
Records—Clipperton Island: on rocks, south of landing
place (Holotype); east side, reef flat (both USNM),.
Littorina sundaica Altena, 1945
(Pl. 349, figs. 1, 2)
Range—Java, Indonesia.
Remarks—The “Sunda” littorine is an ap-
parently quite distinct species with a rather nar-
row geographic distribution according to avail-
able records, having been reported only from
western Java. Due to its small size, however, it
may be overlooked easily and when more collec-
tions are made in nearby areas of Indonesia, the
species may be found more widespread. The
dark-brown color, few spirally incised lines of
sculpture and single white spiral color band visi-
ble internally and externally, together with its
rather narrow shell outline are about the only
distinctive characteristics of this species, yet they
serve to distinguish it from any other known
littorine. The shell of L. sundaica bears a slight
similarity to the species assigned to Rissolittorina
Ponder, 1966, R. alta Powell, and mariae Ten-
ison-Woods, especially due to the spiral white
band on the base of the shell of all three. How-
ever, Rissolittorina appears to be a cool water
group whereas L. sundaica obviously is a tropical
species.
Habitat—Not definitely known, but probably
on shore rocks.
Description—Shell reaching 8.5 mm. (about
5/16 inch) in length, elongate-narrow-turbinate in
shape, average obesity about .55 (4 specimens
range from .53 to .57); shell moderately thick in
structure for its size, imperforate, smooth, except
for a few spirally incised lines on body whorl and
fine growth lines. Overall color of shell nearest to
dark reddish brown (ISCC-NBS color number 44)
becoming somewhat lighter brown in a diffuse
band near suture and aperture edge; with a nar-
row, white spiral color band on base below per-
iphery also showing in aperture. Base hardly
flattened, body whorl subcarinate at periphery.
Whorls 5-7, very slightly rounded. Spire more
than half length of shell, convex, produced at an
angle of about 42°. Aperture subquadrate; outer
lip moderately thick, inner lip nearly straight to
only slightly concave, glazed with a brown callus,
slightly pinched at junction with outer lip. Suture
distinct; whorls may be slightly shouldered.
Sculpture consists of 2-3 weak incised spiral
striae just above periphery and 1-3 below it on
base; fine axial and spiral growth lines are visible
microscopically. Periostracum not evident. Oper-
culum corneus, paucispiral. Nuclear whorls de-
collate in all specimens examined. Radula, animal
and reproduction unknown.
Measurements (mm.)—
length width no. whorls locality
8.5 4.5 7+ (holotype) Tjilaoet
Eureun, Java
2 3.8 5+ “Java”
6.0 3.4 5+ Kaledjetan, Bantam, Java
5.5 3.0 6+ “Java”
Synonymy—
1945 Littorina (Melaraphe) sundaica Altena, Zoologische
Mededeelingen, Leiden, vol. 25, p. 151, fig 2, (Tji-
laoet Eureun, south coast of Java); holotype in Rijks-
museum van Natuurlijke Historie, Leiden, no.
52038.
Records—JAVA: (MCZ, USNM); Kaledjetan, Bantam
(USNM); Tjilaoet Eureun, (Holotype, RNHL).
[05-312]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 451
Littorina acutispira E. A. Smith, 1892
(PI. 349, fig. 3; pl. 351, fig. 1)
Range—Known only from coasts of southern
Queensland and New South Wales, Australia.
Remarks—This and the preceding species, L.
sundaica Altena, are placed provisionally in the
a
aa
ads
Plate 349. Figs. 1,2. Littorina (Melarhaphe) sundaica Altena.
Holotype, from Tjilaoet Eureun, S. coast of Java (Leiden
Museum no. 52038, 8.4 x 4.4 mm.).
Fig. 3. Littorina acutispira E. A. Smith. Lectotype from
Port Jackson, New South Wales, Australia (BM(NH)
91.11.6.216-225, about 7 x 4 mm.).
Figs. 4,5. Littorina lucida Yokoyama. Unique holotype,
from Pliocene of Japan (Geological Institute, Imperial
University of Tokyo, 5 x 3 mm.).
Figs. 6,7. Littorina kozaiensis Nomura and Onisi. Unique
holotype, from Lower Miocene of Japan (Saito H6-on Kai
Museum, Sendai, Japan, Reg. no. 21762, 11 x 8 mm.).
Figs, 8,9. Littorina adonis Yokoyama. Unique holotype,
from Pliocene of Japan (Geological Institute, Imperial
University of Tokyo, 6 x 4 mm.).
subgenus Littoraria. More definite placement
may be possible upon examination of fresh ana-
tomical material, which has not been available for
the present study. The general appearance of L.
acutispira is littorinoid, but certain features, such
as a tendency to form an umbilicus and the
yellow bordered aperture which is dark in its
interior, resemble some of the Fossaridae, notably
F. atratus C. B. Adams of the tropical eastern
Pacific. Because of the generalized nature of Lit-
torina, it is not surprising that some species should
resemble more generalized species of other fami-
lies. The light-colored apertural edge is also seen
in L. africana and there is a tendency toward this
character in L. unifasciata.
Habitat—‘Common in rock pools” (Smith,
1892).
Description—Shell reaching 7 mm. (about 0.3
inch) in length, elongate-turbinate in shape, aver-
age obesity about .57 (7 specimens range from .51
- .63); moderately thick in structure for its size,
tending to become umbilicate, sculptured with
weak spiral striae and irregular, coarse axial lines
of growth. External color variable, from light
yellowish brown to dark brown, usually with a
wide spiral band of brown to gray encircling
body whorl and anterior 2/3 of spire whorls;
speckled on the body whorl with whitish spots.
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Plate 350. Geographical distribution of the species, Littorina
(Littoraria) sundaica Altena, in Indonesia, and of L. acutis-
pira E. A. Smith, in eastern Australia.
[05-313]
452 Littoraria
Aperture dark reddish-brown within and on colu-
mellar lip; outer lip with a broad white band and
a white stripe revolving into it. Base slightly
flattened; body whorl weakly carinate. Whorls
4-6, rounded. Spire more than half the length of
shell, convex, produced at an angle of from 45-
50°. Aperture oval; outer lip moderately thick,
inner lip slightly concave, glazed with a brown
callus, flattened, and pinched near junction with
outer lip; when umbilicus is present there is a
trough just outside columellar callus. Suture dis-
tinct, whorls hardly shouldered. Spiral sculpture
consists of well-spaced, weak spiral striae over
surface of whorls; axial sculpture of irregular
axial lines of growth. Periostracum not evident.
Operculum corneus, paucispiral. Nuclear whorls
decollate in all specimens examined. Radula, ani-
mal and reproduction unknown.
Measurements (mm)—
length width no. whorls locality
7 3.5 6 lectotype
5.6 3.4 5 All from Port Jackson,
5.6 3.1 5 New South Wales,
5.0 2.6 5 Australia
4.5 2.5 5+
4.3 2.7 4+
3.8 2.2, 4
Synonymy—
1892 Littorina acutispira E. A. Smith, Proceedings of the
Zoological Society of London, part 4, (1891), p. 487,
pl. 40, fig 3 (rock pools, Green Point, Watson Bay,
Port Jackson, New South Wales); lectotype BM(NH)
91.11.6.216-225.
Types—One of the 14 syntypes of L. acutispira
Smith (BM(NH) 91.11.6.216-225) is here chosen
as lectotype for the species (see our pl. 349 fig.
of 7 X 3.0 mim,),
Records—AUSTRALIA: Gladstone; Caloundra, both
Queensland (both AMS); Port Macquarie; Catherine Hill Bay;
Middle Harbor (all AMS); Port Jackson, all New South Wales
(AMS; MCZ; USNM).
Littorina infans E, A. Smith, 1892
(Pl. 351, fig. 2)
Range—New South Wales and Queensland,
Australia.
Descriptive Remarks—Littorina infans is in-
cluded provisionally here although its small size
and the lack of any real proof of anatomical
relationships causes me to consider it more a
doubtful species of Littorinidae. Final generic
and possibly even family assignment must await
further investigation.
Joseph Rosewater
Littorinidae
Plate 351. Fig. 1. Littorina acutispira E. A. Smith, from Port
Jackson, New South Wales, Australia (from original figure
in Proc. Zool. Soc., London, for 1891, pl. 40, fig. 3; about 7
x 3.5 mm.).
Fig. 2. Littorina infans E. A. Smith, from Port Jackson, New
South Wales, Australia (from original figure in Proc. Zool.
Soc., London, for 1891, pl. 40, fig. 4; about 3.5 x 2.3 mm.).
Synonymy—
1892 Littorina infans E. A. Smith. Proceedings of the Zoologi-
cal Society of London (1891) p. 488, pl. 40, fig. 4
(Green Point, Watson Bay, Port Jackson [New South
Wales, Australia] “found in rocky pools washed by
ordinary high tides” (Brazier); syntypes BM(NH);
91.11.6.226-235; original measurements 3.5 x 2.3
mm.
Records—AUSTRALIA: NEW SOUTH WALES: Green
Point, Watson's Bay, Port Jackson (Syntypes: BM(NH); AMS);
Sussex Haven; Little Coogee Bay; Shark Island. QUEENS-
LAND: Burleigh Heads; Caloundra; Marouchydore (all
AMS).
[05-314]
November 30, 1970
Littorina kozaiensis Nomura and Onisi, 1940
(P1. 349, figs. 6, 7)
Range—Lower Miocene of Japan.
Remarks—This species was described as re-
sembling L. adonis Yokoyama (see below), but as
having a larger number of spiral grooves. The
unique holotype (pl. 349, figs. 6, 7, a copy of the
original illustration) offers little basis for com-
parison with Recent species.
Synonymy—
1940 Littorina kozaiensis Nomura and Onisi, Japanese Jour-
nal of Geology and Geography, vol. 17, nos. 3 and 4,
p. 191, pl. 19, fig 6 a,b. (Yosuibori, Simizu, Kozai-
mura, Japan); holotype: Saito Ho-on Kai Museum,
Register No. 21762; 11 x 8 mm.
Littorina adonis Yokoyama, 1927
(Pl. 349, figs. 8, 9)
Range—Pliocene of Japan.
Remarks—Placed_ provisionally here in the
subgenus Littoraria, this species resembles L.
undulata, although the strong spiral sculpture is
also reminiscent of Littorinopsis, i.e. L. scabra,
etc, Unfortunately the outer lip and a portion of
the body whorl of the type (pl. 349, figs. 8, 9) are
missing and it is difficult to be sure of their exact
shape.
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 453
Synonymy—
1927 Littorina adonis Yokoyama, Journal of the Faculty of
Science Imperial University of Tokyo, section 2, vol.
1, part 10, p. 451, pl. 51, fig. 8, (Upper Musashino,
Koyasu southern Musashi, Japan); (unique holotype
in collection of Geological Institute Imperial Univer-
sity of Tokyo: 6 x 4 mm.).
Littorina lucida Yokoyama, 1927
(Pl. 349, figs. 4, 5)
Range—Pliocene of Japan.
Remarks—This species is from the same de-
posit as L. adonis but lacks the deeply incised
spiral sculpture. The type of lucida, although of
approximately the same size as adonis is more
slender. This type (pl. 349, figs. 4, 5) is also
broken, but the columella is complete and the
relationship with members of the subgenus Lit-
toraria is more easily confirmed than was the case
with L. adonis.
Synonymy—
1927 Littorina lucida Yokoyama, Journal of the Faculty of
Science Imperial University of Tokyo, section 2, vol.
1, part 10, p. 451, pl. 51, fig. 9 (Upper Musashino,
Koyasu southern Musashi, Japan; unique holotype in
collection of Geological Institute Imperial University
of Tokyo: 5 x 3 mm.).
[05-317]
454 Littoraria
Joseph Rosewater
Littorinidae
[ These occasional blank areas occur between
genera and subgenera to permit the insertion
of new material and future sections in their
proper systematic sequence. ]
[05-318]
November 30, 1970
Subgenus Littorinopsis Morch, 1876
Type: Littorina angulifera (Lamarck, 1822)
The subgenus Littorinopsis is a small group of
mostly tropical species, typified by L. angulifera,
which are usually ovoviviparous and which show
a habitat preference for mangroves or shore vege-
tation. Individual populations of some species,
such as L. scabra and its subspecies L. angulifera,
exhibit a wide range of variation probably due to
the geographical isolation imposed by their meth-
od of reproduction which tends to restrict gene
flow. Nevertheless, within their ranges, these
species are widespread possibly because their
habitat offers opportunities for rafting of adults.
Shells of species of Littorinopsis show strong
development of spiral sculpture which in some
may form multiple carinae (L. carinifera, L.
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 455
scabra) and there appears to be a tendency for
dark coloration and axial color banding and spot-
ting to become strongly developed also. In males
the penis is well-developed, with an open but
folded sperm duct, a thickened base and a nar-
rower distal filament.
There are three Recent and two Tertiary fossil
species of Littorinopsis included here. The ex-
tremely variable L. scabra forms some local pop-
ulations which have been considered as species.
Most of these appear to be only minor genetical or
ecotypical variants, not worthy of even sub-
specific distinction (see scabra synonymy). Two
species closely related to L. scabra (L. carinifera
and L. melanostoma) appear to have developed
sufficient genetic constancy to be considered as
separate species.
Synonymy
1876 Littorinopsis Morch, Malakozoologische Blitter, vol. 23,
p. 135 (type-species by original designation Littorina
subangulata Lamarck [lapsus for angulifera]).
1887 Lamellilitorina Tryon, Manual of Conchology, vol. 9,
pp. 230, 253 (type-species by subsequent designa-
tion, Wenz, 1939: Littorina (Lamellilitorina) albicans
Metcalfe [= L. scabra Linné]}).
Key to Recent Species of Indo-Pacific Littorinopsis
ie olemells lig Diack. 4, sono aad es eee ee Se Sew e nwa eRe melanostoma
hy Columetla lip mot Blak ~ ssid yok Gee a exceed odd Rb he AG embod we Sow Hie PRE A aes 2
2. Two broad white bands revolving within aperture, shell carinate and with
narrow fed axial lipes.OrspOts: 454 4. ha ceo hor 64K SSS Owe e TER carinifera
2. Aperture with many fine lines and spotted with brown inside and out ....... scabra
[05-329 ]
456 _Littorinopsis
Joseph Rosewater
Littorina scabra scabra (Linne, 1758)
(Pls. 325, 352, 353)
Range—Pan-Indo-Pacific, from South Africa to
southeastern Polynesia.
Remarks—The “rough” Littorina, which is one
meaning of the Latin name _ scabra (another is
“scabby,” which would also fit a brown-spotted
shell) is an extremely ubiquitous species through-
out the Indo-Pacific wherever there are man-
groves or even wood pilings. Its comparatively
large size would appear to make it easily visible
were it not well camouflaged on mangrove trunks
and branches. The relatively long synonymy is a
partial indication of its variability. The species
produces an almost endless variety of color and
sculptural forms from the “typical” dark brown
mosaicly banded scabra to the whitish rather
smooth albicans described by Metcalfe. The mor-
phological diversity is probably due to the fact
that scabra is ovoviviparous and produces closely
related, inbred populations within which varia-
tions or mutations may develop and are perpetu-
ated. The same phenomenon takes place in the
Atlantic subspecies, angulifera whose biology is
similar to scabra (Lenderking, 1954; Struhsaker,
1966).
Plate 352. Littorina scabra (Linné). Showing a range of shell
variations.
Fig. 1. Helix scabra Linné (Lectotype figure, from Chem-
nitz, vol. 11, pl. 210, fig. 2074, about 35 x 21 mm.).
Fig. 2. Littorina arboricola Reeve (Lectotype, from Sin-
gapore; BM(NH) 1968321, 31.9 x 25.4 mm.).
Fig. 3. Littorina fortunei Reeve (Lectotype, from China;
BM(NH) 1968309, 16.2 x 7.6 mm.).
Fig. 4. Litorina flammea Philippi (Lectotype, from China;
BM(NH) 1968310, 16.8 x 7.7 mm.).
Fig. 5. Littorina albicans Metcalfe (Lectotype, from Sara-
wak, Borneo; BM(NH) 1968355, 18.8 x 10.8 mm.).
Fig. 6. Litorina scabra articulata Philippi (Lectotype, from
Mindanao; BM(NH) 1968354, 31.5 x 16 mm.).
Fig. 7. Littorina cingulata Philippi (Lectotype, from “north
coast of Australia”; BM(NH) 1968352, 19.4 x 10.7 mm.).
Fig. 8. Littorina intermedia Philippi (Lectotype, from Ne-
gros Id., Philippines; BM(NH) 1968353, 14.2 x 7.8 mm.).
Fig. 9. Littorina luteola Quoy and Gaimard (Lectotype,
from Port Jackson, Sydney, Australia; MHNP, 16.3 x 9
mm.).
Fig. 10. Littorina newcombi Reeve (Lectotype, from Ha-
waii; BM(NH) 1968308, 25.3 x 15.1 mm.).
Fig. 11. Littorina novaehiberniae Lesson (Lectotype, from
Port Praslin, New Ireland; MHNG, 24 x 13.8 mm.).
Fig. 12 Littorina pallescens Philippi (Lectotype, from Min-
danao, Philippines; BM(NH) 1968277, 22.1 x 12.9 mm.).
Fig. 13. Littorina philippiana Reeve (Lectotype, from
“Philippines”; BM(NH) 1968307, 29.6 x 16 mm.).
Figs. 14,15. Littorina scabra rhodea Biggs (Holotype, from
Bundar Abbas, Persian Gulf; BM(NH) 1958.6.13.23, ca.
11.5 x 7.5 mm.).
Littorinidae
Variations in L. scabra generally take the form
of yellow and orange color mutants. It may also
exhibit normal closely-spaced spiral sculpture to
more widely-spaced carinae. These variations oc-
cur throughout the geographic range of the
species more or less randomly. One variation
which appears to be more geographically limited
occurs on the north coast of Western Australia
and was named L. sulculosa Philippi (see pl. 325,
fig. 3). Specimens resembling this form were
collected from ground rocks in a mangrove
swamp on Barrow Island, Western Australia in
1966, and are considered to be ecophenotypes of
L. scabra, having a habitat different from the
latter. Since forms intermediate between scabra
and sulculosa were found at the same locality,
sulculosa does not appear to have validity as a
specific entity.
Differences between L. scabra scabra and its
Atlantic subspecies L. scabra angulifera are dif-
ficult to enumerate as they are mostly qualitative:
angulifera tends to be lighter in color overall,
ranging more toward yellow and orange than the
darker brown scabra; its markings are more dif-
fuse than in most of the latter; the aperture of
angulifera is narrower, more subquadrate and the
columella tends to have a violet hue, while in
Fig. 16. Littorina sieboldii Philippi (Holotype, from Japan;
BM(NH) 1968278, 29 x 15.4 mm.).
Figs. 17,18. Littorina sulculosa Philippi (Lectotype, from
“north coast of Australia”; BM(NH) 1968279, 17.7 x 9.9
mm.).
Fig. 19. Specimen from Arue, Tahiti (USNM 668338, 15.2
x 9.7 mm.).
Fig. 20. Specimen from Mbweni, Zanzibar (USNM 604470,
15 x 7.3 mm.).
Fig. 21. Specimen from near Mombasa, Kenya (USNM
215230, 19.9 x 10.5 mm.).
Figs. 22,23. Specimen from Proserpine, N. Queensland,
Australia (USNM 679498, fig. 22: 23 x 13.2 mm.; fig. 23:
23.4 x 13.2 mm.).
Figs. 24,25. Specimens from Po Bui Id., Sandakan, North
Borneo (USNM 233252, fig. 24: 20.8 «x 11.5 mm., fig. 25:
22.6 x 11.5 mm.).
Figs. 26,27. Specimen from Barrow Id., Western Australia
(USNM 691687, 16.1 x 9.7 mm.).
Figs. 28,29. Specimen from Sekudu Id., Strait of Johore,
Malaysia (USNM 660732, 10 x 6.1 mm.).
Fig. 30. Specimen from Broome, Western Australia (USNM
637333, 20.2 x 11.6 mm.).
Fig. 31. Specimen from Santubong, Sarawak, Borneo
(USNM 671209, 15.1 x 8 mm.).
Fig. 32. Specimen from Guam, Marianas (USNM 426452,
39.6 x 22.7 mm.).
Fig. 33. Specimen from Flat Top Id., near Mackay, Queens-
land, Australia (USNM 622988, 25.4 x 14.2 mm.).
Fig. 34. Specimen from Proserpine, Queensland, Australia
(USNM 679503, 24 x 12.4 mm.).
[05-330]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina
Plate 352. Littorina scabra (Linné). Explanations on opposite page.
[05-331]
458 Littorinopsis
Joseph Rosewater
Littorinidae
scabra the aperture is more broadly rounded and
the columella whitish or stained with brown; in
angulifera there is often a series of brown spots
just within the edge of the outer lip, while in
scabra the entire interior of the aperture shows
the external color, although this is rather variable
in both species. Sculptural characters are essen-
tially similar and isolated specimens are difficult
to distinguish without comparison. I consider the
range of L. scabra to be limited to the tropical
Indo-Pacific, that of angulifera the tropical east
and west Atlantic. Some introductions of L. angu-
lifera into the east Pacific have taken place in the
vicinity of Panama and the notable subspecies L.
scabra abberans (Philippi, 1846), having a dark
brown-rimmed aperture, occurs occasionally in
that area.
A possible explanation for the wide range of
variation in color of L. scabra, in addition to the
fact that populations are more reproductively
isolated than many other marine snails by reason
of being ovoviviparous, is that it is, in effect,
largely a tree snail. Extreme color variation is
recognized within and between populations of
such pulmonate tree snail genera as Polymita and
Liguus. Clench (1968) stated that for tree snails
“coloration is apparently non-selective as there
must be little ocular predation.” When snails
leave the ground and ascend trees, they are im-
mediately free of much of the danger from attacks
by ground-living invertebrates and mammals
which under ordinary conditions may select them
for the familiar subdued coloration usually evi-
denced by many exposed land, freshwater and
marine snails. It may be theorized, therefore, that
in L. scabra color variation is not under the
control of selective forces usually exerted upon
other species of Littorinidae and is, therefore,
freely expressed in many of its populations.
Habitat—Usually found on the trunks and
branches of mangroves and on the ground; may
be present on dock pilings and on sea walls; not
commonly found on shore rocks except in areas
from which mangroves are absent.
Description—Shell reaching 43 mm. (about
1"/16 inches) in length [occasionally larger, but
often smaller], high-turbinate in shape, average
obesity about .57 (15 specimens range from .50 -
.67); relatively thin in structure, usually im-
perforate, and sculptured with spiral striae be-
tween which on the body whorl are raised, split
spiral cords; body whorl subcarinate to carinate at
periphery. Color and color pattern variable, typi-
cally a mosaic of brown blotches arranged axially
or zigzag or in no apparent pattern; occasional
populations with individuals partly or entirely
lacking brown pattern and ranging in color from
reddish orange through medium brown to yellow
and white. Base often moderately flattened, sep-
arated from upper part of body whorl by a moder-
ately to well developed keel at periphery. Whorls
6-9, well rounded. Ratios of aperture and spire
length to length of shell varying from equal to one
exceeding the other. Spire convex, produced at an
angle of from 43-57° depending on population.
Aperture large, rather broadly rounded in outline;
outer lip rather thin even in mature individuals,
curving out widely; inner lip usually concave
posteriorly (above) slightly convex anteriorly and
almost forming a tooth just above where it joins
outer lip near base of columella. Apertural por-
tion of columella heavy, usually white or with
violet to brown stains. Interior of aperture show-
ing same mosaic color pattern as exterior. Suture
distinctly impressed, whorls often showing a
weak but differentiated subsutural cord standing
out from preceding whorl. Sculpture consists of
from 10-12 weak to relatively strong spiral cords
with intervening shallow spiral striae; cords may
or may not be split by secondary striae. Under
magnification close-spaced, fine, wavy spiral
threads are present over entire surface. Axial
sculpture consists of fine lines of growth; occa-
sionally growth lines are well-developed and
some specimens may have spaced axial “ribs;” in
some specimens every other spiral cord is sup-
pressed and shell develops strongly carinate ap-
pearance. Operculum large, thin, corneous, pau-
cispiral. Periostracum not evident. Nuclear
whorls partially decollate in all specimens ex-
amined; remaining portions light-brown, spirally
sculptured, and appear ornate. Radula littorinoid
(2-1-1-1-2). Animal large, darkly pigmented on
surfaces of tentacles, snout and foot; ctenidium
well-developed in both males and females, serv-
ing in latter as partitioned “brood pouch,” oc-
cupying most of dorsal internal surface of mantle
cavity. Penis with greatly thickened base, having
lateral thickened appendage with bulbous ex-
tremity; penis becoming considerably narrower at
tip. Sperm groove open but folded. Reproduction
ovoviviparous; eggs released into mantle cavity
where held in ctenidial “brood pouch;” usually
released in late veliger stage; free-swimming lar-
val life probably very short. Egg size about 0.11
mm.
[05-332]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 459
Plate 353. Littorina scabra (scabra (Linné).
Fig. A. Radula of specimen from Sanga Sanga Id, Sulu
Archipelago, Philippines, ANSP 230667; upper scale is
0.1 mm.
Figs. B,C. Penis of specimen from Eniwetok, Marshall
Ids.; fig. B. view from posterior showing sperm duct
(detail indicates deep duct) fig. C. anterior view; specimen
relaxed with Propylene Phenoxytol; scale is 5 mm.
Measurements (mm.)—
length width no. whorls locality
43.3 25.3 6 Luzon, Philippines
42.1 23.4 7 Oahu, Hawaii
OOet 22.6 7 Guam
38.2 21.4 8 Barrow Id.,
W. Australia
35.3 19.9 8 Barrow Id.,
W. Australia
33.7 18.0 a Luzon, Philippines
30.7 18.5 7 Buka Id., Gulf
of Tomini, Celebes
27.4 16.4 7 Mauritius
24.1 14.4 Jordan River, Guimaras
Philippines
23.6 11.8 8+ Formosa
20.6 11.6 8 Koh Tau, Thailand
19.8 12.1 7+ Formosa
17.4 10.0 7 Vengurla, India
14.1 7.8 7+ Changanoue Strait,
Mombasa, Kenya
11.0 6.0 "i Buena Vista, Guimaras
Id., Philippines
Synonymy—
[1705 Buccinum foliorum Rumphius, d’Amboinische Rariteit-
kamer, p. 98, pl. 29, species Y; prelinnean.]
1758 Helix scabra Linne, Systema Naturae, ed., 10, vol. 1, p.
770; type locality here designated: Amboina, Moluc-
cas; Lectotype here designated: specimen figured by
Chemnitz, vol. 11, pl. 210, fig. 2074.
1791 Buccinum lineatum Gmelin, Systema Naturae ed. 13,
vol. 1, part 6, p. 3493; refers to Knorr, Vergn, 3, pl. 14,
fig. 4, here designated lectotype (no locality given).
1831 Littorina novaehiberniae Lesson, Voyage Autour du
Monde, Coquille, vol. 2, part 1, p. 348 (Port Praslin,
Nouvelle-Irlande [Solomon Islands]. Lectotype here
designated, specimen ex. Lesson in MHNG: 24 x 13.8
mm).
1832 Littorina luteola Quoy and Gaimard, Voyage de |’Astro-
labe, vol. 2, p. 477, pl. 33, figs 4-7 (Port Jackson,
[near] Sydney [Australia]); lectotype here designated
one of 7 syntypes in MHNP: 16.3 x 9.0 mm.
1846 Littorina intermedia Philippi, Proceedings of the Zoo-
logical Society of London, part 13 (1845), p. 141 (here
restricted to Jimamailan, Negros Id., Philippines);
lectotype here selected BM(NH) 1968353: 14.2 x 7.8
mm.; 1847; Abbildungen und Beschreibungen Con-
chylien, vol. 2, p. 223, Litorina pl. 5 fig. 8, (lectotype).
1846 Littorina intermedia punctata Philippi, ibid., (no locality
given in 1846); 1847, ibid; Litorina pl. 5, fig 11
(Tahiti; Elizabeth Island; Natal; Red Sea).
1846 Littorina intermedia articulata Philippi, ibid., (Swan
Point [Western Australia]); 1847, ibid.
1846 Littorina intermedia strigata Philippi, ibid., (Ji-
mamailan, Negros Id. Philippines); 1847, ibid.,
Litorina pl. 5, figs 8-10. (Mergui).
1846 Littorina pallescens Philippi, ibid., p. 142 (Province of
Cagayan de Misamis, Mindanao [Philippines]); lec-
totype BM (NH) 1968277: 22.1 x 12.9 mm.; 1847,
ibid., vol. 3, p. 10 [corrected page] Litorina pl. 6, fig 4.
1846 Littorina sieboldii Philippi, ibid., (Japonia); holotype in
BM(NH) 1968278: 29 x 15.4 mm.; 1847 ibid., p. 9,
Litorina pl. 6, fig 3.
1846 Littorina cingulata Philippi, ibid., (ad oram borealem
Novae Hollandiae); lectotype BM(NH) 1968352: 19.4
x 10.7 mm.; 1847, ibid., p. 11, Litorina pl. 6, fig 5.
1846 Littorina sulculosa Philippi, ibid.; (in Ora boreali Novae
Hollandiae); lectotype BM (NH) 1968279: 17.7 x 9.9
mm.; 1847, ibid., p. 18, Litorina, pl. 6, fig 10.
1847 Litorina scabra flammulata Philippi, Abbildungen und
Beschreibungen Conchylien, vol. 2, p. 222, Litorina
(Panay; Singapore)
1847 Litorina scabra articulata “Menke” Philippi, ibid.;
pl. 5, fig. 4, (Mindanao here restricted); lectotype
BM(NH)1968354: 31.5 x 16 mm.)
1847 Litorina scabra punctata Philippi, ibid., pl. 5, fig 5.
(Masbate; Singapore).
1847 Litorina scabra suturalis Philippi, ibid., pl. 5, fig 7
(Kanguruh-Inseln, dem Meersbusen Georgs des Vier-
ten; Canton)
1847 Litorina scabra lutea Philippi, ibid., pl. 5, fig. 6 (Mas-
bate; Philippines, Canton)
1847 Litorina scabra rubra Philippi, ibid., refers to Chemnitz
[vol. 11, pl. 210] fig. 2075 (Mindoro, Philippines;
Canton)
1847 Litorina scabra ventricosa Philippi, ibid., pl. 5, fig 8
(China; Mindanao; Tonga Tabu; Neu Irland; Neu
Guinea); not L. ventricosa Philippi, ibid., vol. 3, p.
15.
1847 Litorina fammea Philippi, ibid., vol. 3, p. 16, Litorina,
pl. 6, fig. 21 (China); lectotype BM (NH) 1968310:
16.8 x 7.7 mm.
1847 Litorina sinensis Philippi, ibid., vol. 3, p. 52, Litorina pl.
6, fig 23 (China).
1852 Littorina albicans Metcalfe, Proceedings of the Zoologi-
cal Society of London, part 19 (1851) p. 73 (Sarawak,
Borneo); lectotype BM(NH) 1968355: 18.8 x 10.8
mm.; 1857, Reeve, Conchologia Iconica, vol. 10, pl. 9,
figs. 44a, b.
1857 Littorina philippiana Reeve, ibid., pl. 5, figs 22 a,b
(Philippine Islands); lectotype BM(NH) 1968307,
29.6 x 16 mm.
1857 Littorina arboricola Reeve, ibid., pl. 6, fig 27 a,b (Sin-
gapore); lectotype BM(NH) 1968321: 31.9 x 25.4 mm.
1857 Littorina newcombi Reeve, ibid., pl. 6, fig 28 a,b (Sand-
wich Islands); lectotype BM (NH) 1968308: 25.3 x
15.1 mm.
1857 Littorina fortunei Reeve, ibid., pl. 9, figs 42 a,b China);
lectotvype BM(NH) 1968309; 16.2 x 7.6 mm.
[05-333]
460 Littorinopsis
Joseph Rosewater
Littorinidae
1871 Litorina strigata Lischke, Malakozoologische Blatter,
vol. 18, p. 148 (Nagasaki [Japan]); type: Academy
Science, Leningrad?; 1871 Japonische Meeres-
Conchylien, vol. 2, p. 73, pl. 5, fig. 22.
1871 Melaraphe (Littorina) blandfordi Dunker, Malakozoo-
logische Blatter, vol. 18, p. 150 (Rockhampton [Aus-
tralia]); type in Berlin Museum?
1878 Litorina scabra concolor Weinkauff, Systematisches
Conchylien-Cabinet, vol. 2, part 9, p. 37, pl. 4, figs
8-10 [not fig. 11 as indicated in text = L. undulata]
(Indo-Pacific) [in part].
1878 Litorina scabra minor Weinkauff, ibid., pl. 4, figs. 16-18
(Indo-Pacific); refers to L. intermedia Philippi and to
L. newkombi (sic) Reeve.
1878 Littorina newkombi ‘Reeve’ in Weinkauff, ibid., pp. 37,
38 [error for L. newcombi Reeve].
1885 Littorina scabra tenuis Nevill, Hand-List of Mollusca in
the Indian Museum, part 2, p. 146 (Arakan [Burma]);
type in Indian Museum, Calcutta?; not L. tenuis
Philippi, 1846. [=L. undulata].
1885 Littorina pallescens? erronea Nevill, ibid., p. 148 (Sin-
gapore); type in Indian Museum, Calcutta?
1885 Littorina filosa subcingulata Nevill, ibid., p. 149 (Port
Jackson [Australia]); type in Indian Museum, Calcut-
taP
1885 Littorina conica delicatula Nevill, ibid., p. 149 (Port
Canning and False Point (Bengal); type in Indian
Museum, Calcutta?)
1885 Leptopoma (?) ardouinianum Heude, Memoires con-
cernant I’Histoire naturelle de ]’Empire Chinois,
Cahier 3, Notes sur les Mollusques terrestres de la
vallee du Fleuve Bleu, p. 95, pl. 25. figs. 8, 8a (ad
rupes Tonquinenses in portu dictu A-long)
1900 Littorina philippina von Martens, Biologia Centrali-
Americana, Land and Freshwater Mollusca, p. 584;
emendation for L. philippiana Reeve, 1857.
1958 Littorina (Melaraphe) scabra rhodea Biggs, Journal of
Conchology, vol. 24, no. 8, p. 272 (Bundar Abbas
[Persian Gulf]); holotype BM(NH) 1958.6.13.23, ca.
11.5 x 7.5 mm.)
Types—In the original description of Helix
scabra Linné refers to the Museum Ludovicae
Ulricae, leading one to believe that a specimen
under that name was present in that collection.
Such a specimen is not contained, however, in the
“MLU” of Odhner’s unpublished list [1953], nor
is H. scabra represented in the Linnean Shell
Collection in London. Furthermore, in the 12th
Edition Linné did not refer to the Ulricae collec-
tion under scabra. In order to clarify the concept
of H. scabra, one of the figures referred to by
Hanley (1855), is here designated as lectotype:
Chemnitz, Conchylien Cabinet, vol. 11, pl. 210,
fig. 2074 (see pl. 352, fig. 1). The specimen
represented in that figure may be in the Zoologi-
cal Museum, Copenhagen. The type locality for
H. scabra is here designated as Amboina, Moluc-
cas, as no locality was given by Linné.
The location of holotypes and lectotypes of the
other synonyms of scabra are mentioned in the
synonymy where this information is known to
me. Types of species described by Philippi in the
“Abbildungen” are probably in the Berlin Muse-
um except in those few cases where I designated
as lectotypes the figured specimens found in the
British Museum (NH); see Synonymy.
The lectotype of L. albicans Metcalfe, men-
tioned in the synonymy, which is the specimen
figured by Reeve, is undoubtedly part of the
original syntypic series, as Metcalfe’s collection
was purchased by Reeve and parts of it were
resold (The Athenaeum, No. 1906, May 7, 1864, p.
630).
Records—SOUTH AFRICA: Port Alfred, nr. Grahamstown,
Bathurst District, Cape Province (USNM, MCZ):; Durban;
Isipingo, both Natal (both MCZ). MOZAMBIQUE: Inhaca
Island, Delagoa Bay (NMW; ANSP). TANZANIA: Dar es
Salaam; 1.5 mi. N.W. of Magogoni; Kendwa Island; Kun-
duchi; Bagamoya; Tanga (all MCZ); Mboamagi, 9 mi. S. of
Dar es Salaam (USNM); 2 mi. S. by W. of Chwaka, East
Zanzibar; Bungi, Kiwani Bay, $.W. Zanzibar; 1 mi. N. of
Chukwani, W. Zanzibar (all ANSP); Mbweni, 4 mi. S. of
Zanzibar City; Bweju, S.E. Zanzibar (both USNM). KENYA:
Mombasa Island (ANSP); Straits at Changanoue, 3.75 mi.
from Mombasa, Kenya (USNM). MADAGASCAR: S. of
Anivorano, 12 mi. N. of Ambodifototra, W. coast of Ile Ste.
Marie; Ambodifotgtra, at causeway to Ilot Mandane, Ile Ste.
Marie; S.W. shore Ie des Nattes, S. of Ile Ste. Marie; Soalary,
16 mi. S. of Tuléar; nr. Grottes de Sarodrano, 10 mi. S.W. of
Tuléar; Tuléar (all MCZ); Nossi Be (ANSP; MCZ). SEY-
CHELLES: Aldabra Island (Y.P.M.; USNM); Menai Island,
Cosmoledo Atoll; Anse a la Mouche, Mahé Island; Northwest
of Moyenne Island (all YPM). MAURITIUS: Point d’Espy, 1
mi. N. by E. of Poste de Flacq (USNM, ANSP); 3/4 mi. N. of
Black River Bay (ANSP). MALDIVES: Male Harbor, Male
Atoll; between Mafilefuri and Maro Islands, Fadiffolu Atoll
(both ANSP); Gan, Addu Atoll (YPM; ANSP). INDIA: Bom-
bay (USNM; MCZ): Vengurla, N. of Goa; Goa; Khumpta
(Kumta), N. of Kanaru; Netravati River, Mangalore; Tuticorin;
Adyar River estuary, Madras; Port Canning (all USNM);
Cochin Harbor, Kerala; Rameswaren Island, Pamban, Palk
Strait (both ANSP); Calcutta (MCZ). CEYLON: (MCZ,
ANSP). BURMA: Victoria Point (MCZ). THAILAND: Ko
Contee, Ranong (MCZ); Ko Sindarar Nua (Chance Island); Ko
Phi Phi; Pulau Tanga, Butang Group (all USNM); Songkla
(MCZ); Ban Tha Kham; Ko Phluai; Ko Tao (all USNM);
Mutapone Island, Chumpon; Ban Hua Hin; Ko Si Chang, 40
mi. S. of Menam River; Bang Poo, Paknam; Ang Hin, Chol-
buri Province; Bang Saen, Cholburi Province (all MCZ); Ko
Chang; Ko Kut (both USNM). CAMBODIA: Kampot (MCZ).
VIETNAM: Saigon (MCZ; MHNP). MALAYSIA: Penang
(USNM, MCZ):; N.E. corner, Pulau Lumut, Port Swettenham;
Pulau Besar, Malacca Strait, S.E. of town of Malacca; Cape
Rachardo, Straits of Malacca; Sekudu Island, Strait of Johore
(all USNM); Singapore (USNM, ANSP, MCZ). CHINA:
Kiautschou, nr. Tsingtau, Yellow Sea; Spider Island, Fukien
Province; Amoy (all USNM); Hong Kong (BM(NH), USNM,
ANSP); Macao; Hainan (both ANSP. JAPAN: Awa (ANSP,
MCZ). RYUKYU ISLANDS: (USNM). TAIWAN (USNM,
ANSP, MCZ). PHILIPPINE ISLANDS (Many localities; see
map). SUMATRA: Moro Id. (SMF); Pulau Melila, S. of
Udjung Batu, Banyak Islands; S.E. coast of Pulau Nias; Pulau
Bai, Batu Group (all USNM); Padang (MCZ); Pulau Siburu,
N. of Sipora; S.W. tip of Sanding Island, Mentawi Islands
(both USNM); Tapanuli Bay, Sibolga (ZMA). JAVA: Men-
scheneter Island (MCZ, ANSP, USNM); Kaledjitan, Bantem
(USNM); Djakarta (USNM, ZMA). BALI: Koeta Beach
(USNM). BORNEO: Santubong, Sarawak (USNM, MCZ);
Kudat Bay (ANSP, USNM); Tajong Aru, Jesselton (USNM);
W. Marudu Bay (USNM, ANSP); Po Bui Island, Sandakan;
Karamuntig, Sandakan; sandy plain, Sandakan Bay; Bohay-
dulong Islands (all USNM). CELEBES: Dago Bay, Sangihe
Islands; Manado (both MCZ); Likupang; Papajato River;
Bukabuka Island, Gulf of Tomini; Limbe Island, Gulf of
Tomini (all USNM); Wowoni Id., N. of Butung Island (MCZ);
[05-334 ]
November 30, 1970
Labuan Blanda Island, Butung Strait; Labuandata Bay, Gulf
of Boni (Bone) (both USNM). MOLUCCAS: Talaud Islands;
Morotai Island; Ternate; Kahatola Island, Loloda Islands,
N.W. coast of Halmahera; Pajahi Bay, Halmahera Island;
Kasirota Bay, Kasirota Island; Batjan Island; Oong Bay, Man-
dioli Island; Buru Id.; Tengah Island, nr. Buru; Manipa
Island (all MCZ); Amboina (ANSP); Kobroor, Aru Ids. (SMF);
COCOS-KEELING ISLANDS: N.E. end Direction Island
(USNM). AUSTRALIA: QUEENSLAND: Darnley Island
(MCZ); Murray Island (AMS); Thursday Island, all Torres
Strait; mouth of Norman River (both WAM); mud flats nr.
Pilot Station, Karumba; Mapoon all Gulf of Carpentaria; Cape
Sidmouth; Burkitt Island, nr. Port Stewart; Three Isles (all
AMS): Cooktown (AMS, MCZ); Redlynch (MCZ); Cairns
(USNM):; Green Island, off Cairns (USNM, MCZ, AMS);
Atherton (MCZ); Flying Fish Point (AMS); Dunk Island;
Palm Island (both ANSP, AMS); Cape Bowling Green
(USNM): Bowen (AMS, MCZ); Hayman Island (AMS); Shute
Harbor (USNM): Hamilton Island; Lindeman Island (both
AMS); Proserpine (MCZ); Flat Top Island, nr. MacKay
(USNM):; Percy Island, off Palmerston (MCZ); Yeppoon
(AMS): Gladstone (USNM, ANSP); Port Curtis; Port Vernon;
Pialba; Urangan (all AMS); Moreton Bay (MCZ); Sandgate,
nr. Brisbane (ANSP, AMS); Stradbroke Island (WAM; AMS).
NEW SOUTH WALES: nr. Fingal; Broken Bay; Palm Beach
and Careel Bay, Pittwater (all AMS); Sydney (MCZ, AMS,
USNM, MHNP); Botany Bay; Woolaware Bay, nr. Cronulla
(both AMS); Wollongong (ANSP); Jervis Bay, Huskisson;
Narooma, (both AMS). WESTERN AUSTRALIA: Red Bluff,
Shark Bay; Dirk Hartog Island; Herald Bight, N.E. side of
Peron Peninsula, Shark Bay; Denham, Peron Peninsula, Shark
Bay (all WAM); Carnavon (WAM, USNM); 20 mi. N. of
Cardabia; Monte Bellow Islands (both WAM); Barrow Island
(WAM, USNM); Cape Bossut (BPBM); mouth of False Cape
Creek, La Grange Bay (USNM, ANSP); Broome (AMS, MCZ,
USNM, ANSP); Buccaneer Archipelago; Vansittart Bay (both
AMS). NORTHERN TERRITORY: Napier Bay, Melville Id.
(WAM); Darwin (MCZ, USNM, ANSP). NEW GUINEA:
Hollandia; Seleo Island, off Aitape; Milne Bay; Hilimoi
Mission, Milne Bay, Papua (all USNM); Collingwood Bay;
Woodlark Island (Murua Island) (both AMS); Port Moresby
(USNM, AMS, MCZ, BPBM, ANSP); Oriomo River, Papua;
Daru, Papua (both MCZ); Merauke (MCZ, USNM); Fakfak
(MCZ); 4 mi. N.W. of Korido Village, Soepiori, Schouten
Islands (ANSP); Sorendidori Bay, Soepiori, Schouten Islands
(MCZ); Samberbaba, Japen Island; Rainbawi Point, Japen
Island; '/2 mi. E. of Kaipoeri Village, Koeroedoi Island,
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 461
Geelvink Bay (all ANSP); Mios Woendi, Schouten Islands
(USNM): ADMIRALTY ISLANDS: Koruniat Island (ANSP).
BISMARKS: New Britain (AMS); New Ireland (MCZ). SOL-
OMON ISLANDS: Malaita (MCZ; BPBM); Three Sisters
Island; Marava Lagoon, New Georgia (both MCZ); Pavuvu
Island, Russell Ids. (USNM); Florida Island (MCZ, USNM);
Tulagi, Florida Island (MCZ); Guadalcanal (USNM, AMS).
SANTA CRUZ GROUP: Sunday Bay, Vanikoro Island (AMS).
NEW HEBRIDES: Qakea, (AMS); Ver, N.E. Santa Maria Id.,
both Banks Group (USNM); Hog Harbor, Espiritu Santo
Island (MCZ); Eromanga (AMS). NEW CALEDONIA: mouth
of Ponandou River, E. of Touho (ANSP); Houailoo (BPBM);
Ile Nou; Baie du Prony, S. side Mount Dore, nr. Noumea;
Ricaudy Reef, Noumea; la Roche Percée, Bourail (all ANSP);
llot Amédie (AMS). FIJI ISLANDS: Yasawa Islands (USNM);
Savusavu, Vanua Levu (MCZ); N. shore Nananu-i-ra Island,
(MCZ, USNM):; Suva (MCZ, USNM, BPBM); Nadronga
(USNM, MCZ); Mbalavu (BPBM); Korolevu (ANSP); Lautoka
(MCZ); Nadi Airport, all Viti Levu (USNM); Onega Levu;
Navutu-i-loma (both BPBM). MARIANAS: Saipan (ANSP);
Guam (USNM, BPBM). PALAU ISLANDS: Babelthaup Island
(USNM, BPBM, ANSP); Koror Island (BPBM); Malakal Pass,
W. of Aurapushokaru Island; S.E. end of Eil Malk (both
ANSP); Peleliu (BPBM). CAROLINE ISLANDS: Yap Island,
(MCZ, ANSP); Moen Island; Dublon Island, both Truk Is-
lands (both BPBM); Ponape (BPBM); Kusaie Island (USNM,
BPBM); Kapingamarangi (USNM); MARSHALL ISLANDS:
Eniwetok; Arno; Jaluit Atolls (all USNM). GILBERT IS-
LANDS; Abaiang Island (BPBM, ZMA); Kingsmill Group
(ANSP, USNM). HAWAIIAN ISLANDS: Kauai; Oahu;
Molokai and Hawaii Islands (Many localities, see map). LINE
ISLANDS: Palmyra Island (MCZ, BPBM); Fanning Island,
(BPBM). ELLICE ISLANDS: Vaitupu; Nukulailai (both
USNM). WALLIS ISLANDS: btwn, Luaniva and Fungalei
Islands (USNM). TONGA: Velitoa Tongatapu (BPBM).
SAMOA: Toamua (BPBM); Saluafata, both Upolu (ANSP);
Pago Pago (BPBM, USNM, ANSP); Fagaitua Bay, both Tu-
tuila(MCZ). SOCIETY ISLANDS: Fanui Bay (USNM); N. end
Vaitape; between. Pererau Fai and Araara Pupu, District of
Anau, all Bora Bora (both ANSP); Bay Vaiore; around point
from Hipu, both Tahaa; Uturoa; Tevaitoa, all Raiatea; Lac
Maheva; Port du Bourayne, both Huahine; W. fork of the
Opunohu River, Moorea; N.W. of mouth of Nahoata River,
Pirae; 9 km. from Taravao, Vairao, Taiarapu Peninsula; Motu
Fenuaino, Tautire (all USNM); S. side of Faratea Point, Faaa
District, all Tahiti (ANSP). GAMBIER ISLANDS: Rikitea,
Mangareva; Vaituatai Bay, Mangareva, (both BPBM).
20 40° 60 80 \oo° 120 140° (60° 180 160° 140° 120°
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Plate 354. Geographical distribution of Littorina (Littorinop-
sis) scabra scabra (Linné) in the Indo-Pacific faunal region.
[05-335]
462 _Littorinopsis
Littorina melanostoma Gray, 1839
(Pls. 325, 355)
Range—Southeast coast of India to Borneo.
Remarks—The “Black-mouth” littorine is a
very distinctive species within its rather narrow
range in south Asia and the East Indies. Its
graceful, attenuate outline, dark brown columella
Joseph Rosewater
Littorinidae
and color pattern offer a combination of charac-
ters enabling rapid identification and may qualify
this species for the title “most attractive Lit-
torina.” Nearest relatives appear to be L. cari-
nifera Menke, which also has a fairly restricted
distribution in the same general region, and L.
scabra Linné which is pan-tropical. Among
scabra, carinifera and melanostoma, there are
Plate 355. Littorina melanostoma Gray (figs. 1-4) Littorina
carinifera Menke (figs. 5-13).
Figs. 1,2. Lectotype of L. melanostoma Gray, from Pe-
nang, Malaysia (BM(NH) 1968364, 23.8 x 11.6 mm.).
Figs. 3,4. Port Swettenham, Malaysia (USNM 661028, 28.1
x 13.9 mm.).
Fig. 6. Lectotype figure of Phasianella carinifera Menke
(from Philippi, Abbildungen and Beschreibungen Con-
chylien, vol. 2, pl. 5, fig. 22; about 21 x 12 mm.).
Figs. 5,7. Sandakan, North Borneo; note similarities to
lectotype figure (USNM 658105a, 16.5 x 10.9 mm.).
Figs. 8,9. Lectotype of Littorina rubropicta von Martens,
from King Id. Bay, Mergui Archipelago, Burma (BM(NH)
87.3.10.140-144, 18.3 x 10.5 mm.).
Figs. 10,11. Lectotype of Littorina conica Philippi, from
Java (BM(NH) 1968225, 22.4 x 12.7 mm.).
Figs. 12,13. Bombay, India (USNM 90470, 21.6 x 12.4
mm.).
[05-336]
November 30, 1970
many similarities. All three occupy a_ similar
habitat, anatomical details are remarkably simi-
lar, and although spawning behavior has not been
verified for melanostoma and carinifera, they
very probably reproduce ovoviviparously.
Habitat—Estuarine areas on mangrove trees
and grasses.
Description—Shell reaching 28.2 mm. (about 1
inch) in length, attenuately conical in shape;
average obesity about .48 (23 specimens ranged
from .44 to .54); relatively thin but strong in
structure, imperforate, and sculptured with shal-
lowly incised spiral striae, overall microscopic
wavy spiral threads and fine, oblique axial lines
of growth. External coloration rather uniform;
ground color yellowish white, with closely
spaced punctate to elongate-rhomboidal brown
markings located between the spiral striae, usual-
ly arranged axially but sometimes obliquely or in
zigzag pattern; sometimes with white dashes be-
tween brown spots; tip of spire often gray; aper-
ture yellowish white or with outer brown color
markings partially showing through; columella
dark chocolate-brown. Base hardly flattened; only
a very weak keel at periphery. Whorls 6-8, usually
rather flat-sided. Spire usually somewhat more
than half the length of shell, produced at an angle
of 42-44°. Aperture oval; outer lip rather thin,
often narrowly shouldered; inner lip not greatly
thickened, weakly concave, the posterior 2/3
glazed with dark brown callus, white where it
joins outer lip at anterior extremity. Suture weak-
ly impressed. Sculpture consisting of about 6
shallow spiral striae on spire whorls, persisting
onto body whorl where as many as 18 may be
present over length of whorl; area between striae
flattened and each may be divided by single,
weak, secondary stria; entire surface covered by
fine microscopic wavy spiral threads; floor of
striae often appear to be finely, shallowly and
closely punctured. Axial sculpture consisting of
rather regularly spaced oblique growth lines be-
coming coarser near outer lip. Operculum large,
corneous and paucispiral. Periostracum not evi-
dent in specimens examined. Nuclear whorls par-
tially decollate in all specimens examined; re-
maining portions smooth and glassy, light grayish
tan in color; succeeding whorls sculptured spiral-
ly. Radula typically littorinid (2-1-1-1-2); central
tooth broad and low (similar to L. scabra and L.
carinifera).
Animal darkly pigmented on surfaces of ten-
tacles snout and foot. External appearance of
animal removed from shell: in female the “‘cover-
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 463
ing-capsule gland complex” (see Fretter and
Graham, 1962) located at posterior right of cteni-
dium, is very distinct, pigmented, and forms a
spiral pattern; in male verge is greatly thickened
from base to one-half its length; distal portion
filamentous; sperm duct deeply folded on floor of
mantle cavity and along posterior edge of thick-
ened base of verge and inner edge of filamentous
portion. Nothing reported concerning spawning
or development of this species, although it is
suspected that species is ovoviviparous. In both
male and female the ctenidia are greatly enlarged
and occupy most of the area of mantle cavity roof,
darkly pigmented and conspicuous.
Measurements (mm.)—
length width no. whorls locality
28.2 14.1 8 Port Swettenham,
Malaysia
27.8 14.1 8 Chaga, Malaysia
5.1 12 8 Nakhorn Si
Thammarat, Thailand
25.9 12.9 7+ Nakhorn Si
Thammarat, Thailand
24.4 11.5 7 Ko Chang, Thailand
24.0 10.9 7 Port Swettenham,
Malaysia
22.6 11.1 T+ Sarawak, Borneo
22.3 12.0 7 Kranji, Singapore
21.9 9.6 8 Burma
21.1 10.1 8 Laem Ngob, Thailand
20.4 102 6+ Port Canning, India
20.0 9.7 8 Vietnam
19.2 9.4 6+ Taiwan
14.9 ie 5+ Hong Kong
J Ke
_[MELANOSTOmaA] *%
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0° 80° 100° 120° \40°
Plate 356. Geographical distribution of Littorina (Littorinop-
sis melanostoma (Gray) in south Asia and Indonesia.
[05-337]
464 Littorinopsis
Synonymy—
1839 Littorina melanostoma Gray, The Zoology of Captain
Beechey’s Voyage—in His Majesty's Ship Blossom,
Mollusks, pl. 140 (Indian Ocean; [Penang, Malaysia,
here restricted]) lectotvpe B.M. (NH) 1968364: 23.8 x
11.6 mm.; non L. melanostoma A. J. Krynicki, 1837,
Bull. Soc. Imp. des Nat. de Moscow, Ann. 1837, No.
2, p. 60, nomen nudum.
1885 Littorina melanostoma articulata Nevill, Hand List of
Mollusca in the Indian Museum, part 2, p. 151 (Hong
Kong; type in Indian Museum, Calcutta).
Types—Littorina melanostoma apparently is
one of the species not collected during the voyage
of the Blossom as Gray gave for a type locality,
“Indian Ocean” which the expedition did not
reach (Rosewater, 1968). One of the 5 syntypes
from Gray’s collection is here selected as lecto-
type: BM(NH) 1968364, 23.8 x 11.6 mm (pl. 355,
figs. 1, 2). The type locality is here restricted to
Penang, Malaysia.
Records—INDIA: Port Canning (USNM, RNHL); Manali,
off Mandapam (MCZ). CEYLON: (E. von Martens, 1887).
BURMA: King Island Bay, Mergui Ids. (E. von Martens,
1887). THAILAND: Ko Chang (USNM); Ang Hin (MCZ); Ban
Tamru, both Cholburi Province (ZMA); Laem Ngob; Pak
Phun (both USNM); Pak Paying, Nakhorn Si Thammarat
(MCZ). MALAYSIA: N.E. corner of Pulau Lumut, Port Swet-
tenham (USNM); Malacca (MCZ). SINGAPORE: Kranji
(ANSP, USNM). CHINA: Hainan; Tsi Mei, Amoy; Hai-
ch’eng, Fukien Province (all ANSP); Castle Peak Bay, Hong
Kong (NMW). TAIWAN: (USNM). VIETNAM: Saigon River,
Cochin China (USNM, MCZ). INDONESIA: Palau Weh
(RNHL); Telok Niboeng (ZMA); Tandjung Tiram, all Sumatra
(ZMA); Djepara (RNHL); Surabaja, both Java (ZMA; RNHL;
SMF); Santubong, Sarawak, Borneo (MCZ, USNM).
Littorina carinifera (Menke, 1830)
(Pls. 325, 355)
Range—South and southeast Asia from West
Pakistan to the Philippines and Borneo.
Remarks—Of the several forms which have
been described as being closely related to Lit-
torina scabra, only two appear worthy of recogni-
tion as distinct species: Littorina melanostoma
Gray and Littorina carinifera Menke. The latter,
“carinate littorine,’ in its most extreme form,
lives up to its descriptive name often having as
many as 3-5 rather strong carinae on the body
whorl. Variation is considerable, however, and
there are populations in which carinae are present
only at the periphery. The species may be recog-
nized by its rather pyramidal shape, the color
pattern of rather straight to oblique or zigzag
reddish axial bands on a yellowish brown back-
ground, and by the presence, usually, of dark
color bands within the aperture.
Joseph Rosewater
Littorinidae
Habitat—On bushes or low shrubs along shore;
mangrove swamps.
Description—Shell reaching 22.2 mm. (about
0.9 inch) in length, pyramido-conical in shape;
average obesity about .63 (23 specimens range
from .55 to .71); relatively thick in structure,
imperforate, often multiply carinate; otherwise
sculptured with well-marked spiral striae, overall
microscopic wavy spiral threads (often obscured
by wear) and fine, regular, closely spaced axial
lines of growth. External coloration rather uni-
form; ground color yellowish to light-brown,
with axial pattern of nearly straight to oblique
reddish brown lines. In carinate specimens pat-
tern consists of reddish brown spots primarily on
carinae. Aperture grayish white with three or
more dark brown bands revolving within; colu-
mella white. Base somewhat flattened; periphery,
at least, markedly carinate. Whorls 6-7, hardly
rounded—to rather flat sided. Spire usually some-
what more than half the length of shell, produced
at an angle of 46-72° (depending on specimen).
Aperture roundly oval; outer lip rather thin but
sometimes thickened within; inner lip not greatly
thickened, occasionally with a denticulate swel-
ling near its base, white in color. Suture moder-
ately to well impressed in highly carinate speci-
mens. Sculpture consisting of about 10 often
punctate spiral striae on spire whorls, persisting
onto body whorl where as many as 20 may be
present over length of entire whorl; in non-
strongly carinate specimens areas between striae
flattened to moderately well rounded as incipient
carinae; in strongly carinate specimens at least 3
inter-stria areas raised as rather strong carinae and
others may be moderately raised; in these speci-
mens striae are deep furrows and occasionally
may be sculptured intrinsically with closely-
spaced axial wrinkles; entire surface covered by
fine microscopic wavy spiral threads, the latter
often obliterated by wear. Axial sculpture consist-
ing of rather regularly-spaced axial growth lines.
Operculum moderate in size, corneous, pauci-
spiral. Periostracum not evident in specimens
examined. Nuclear whorls partially decollate in
all specimens examined; remaining portions
smooth, shining, light tan in color; succeeding
whorls spirally sculptured.
Animal including radula resembling L. scabra
and L. melanostoma; darkly pigmented on upper
surfaces of tentacles, snout and foot. Ctenidial
area darkly pigmented showing through mantle;
in female “covering-capsule gland complex” (see
Fretter and Graham, 1962) located at posterior
[05-338]
November 30, 1970
right of ctenidium is very distinct, its spiral
outline marked with dark pigment; in male verge
greatly thickened at base and for about one-half
its length; thickened portion terminally bulbous;
distal portion filamentous; sperm duct deeply
folded, running along posterior edge of verge.
Nothing reported concerning spawning or devel-
opment, but probably ovoviviparous.
Measurements (mm.)—
length width no. whorls locality
29:9 1252 7 Bombay, India
19.0 112 7 Manapla, Negros Occi-
dental, Philippines
18.6 12 6 Zamboanga, Mindanao,
Philippines
17.8 11.1 7 Iloilo, Panay,
Philippines
17.0 10.6 7 Singapore
16.7 11.4 6 Pancol, Palawan,
Philippines
16.4 11.2 6 Po Bui Id., Sandakan,
North Borneo
14.3 9.0 7 Mangalore, India
13.8 8.4 6+ Silaga R., Samar,
Philippines
10.4 74 4+ Djakarta, Java
Synonymy—
1830 Phasianella carinifera Menke, Synopsis methodica Mol-
luscorum, Pyrmont, edition 2, pp. 51, 141 (locality
unknown).
1832 Littorina perdix King and Broderip, Zoological Journal,
vol. 5 (1831), p. 345 (no locality given),
1846 Littorina conica Philippi, Proceedings of the Zoological
Society of London, part 13 (1845), p. 141 (Java);
lectotype BM(NH) 1968225: 22.4 x 12.7 mm.; 1847,
Abbildungen und Beschreibungen Conchylien, vol.
3, p. 9, Litorina, pl. 6, figs. 1, 2.
1885 Littorina carinifera pyramidalis Nevill, Hand-List Mol-
lusca in Indian Museum, part 2, p. 151 [nomen
nodum |].
1885 Littorina carinifera laevior Nevill, ibid. [nomen nudum ].
1887 Littorina rubropicta von Martens, Journal of The Lin-
nean Society, Zoology, vol. 21, p. 170, pl. 16, figs 2 a-f
(King Island Bay, Mergui Archipelago); lectotype
BM(NH) 87.3.10. 140-144: 18.3 x 10.5 mm.
1932 Litorina carnifera Sherborn, Index Animalium, section
2, part 31, p. 638 [error for L. carinifera].
Types—According to Dance (1966) Menke’s
collection was dispersed and therefore, it is most
unlikely that the original type specimen of L.
carinifera can be located. Fortunately Philippi
(1847, Abbildungen und Beschreibungen Con-
chylien, vol. 2, Litorina, p. 227, pl. 5, fig 22)
apparently had access to Menke’s collection and
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 465
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Plate 357. Geographical distribution of Littorina (Littorinop-
sis) carinifera (Menke) in South Asia, Indonesia and the
Philippines.
figured what he called “the original example”.
That figure is here selected as the lectotype of
Phasianella carinifera Menke (pl. 355, fig. 6). The
type locality is here designated as Negros Oc-
cidental, Philippines, one of the localities men-
tioned by Philippi.
The location of the type-specimen of L. perdix
King and Broderip is unknown. It may be in the
BM(NH) although it was not found during a
recent visit. The lectotypes of L. conica Philippi
and L. rubropicta von Martens are in the BM(NH)
as mentioned in the synonymy (pl. 355 figs. 8-11).
Records—WEST PAKISTAN: China Creek, Karachi
(MCZ). INDIA: Bombay (MCZ, USNM); Vengurla, N. of Goa;
Netravati River, Mangalore (both USNM). BURMA: King
Island Bay, Mergui Ids. (E yon Martens, 1887). THAILAND:
Ko Kut (USNM). MALAYA: N.E. corner Pulau Lumut, Port
Swettenham; Singapore (both USNM). PHILIPPINES:
Medio Island, Galera Bay, Mindoro; Hoilo, Panay; Victorias;
Manapla, both Negros Occidental; W. of Catbalogan; Silaga
River, both Samar; Zamboanga, Mindanao; Pancol, Palawan
(all USNM); Busuanga (RNHL). INDONESIA: Belawan, Su-
matra (RNHL; ZMA); Panahatan, Sumatra; Karanganjar, Java
(both ZMA); Tandjung Priok, Java (RNHL); Djakarta, Java
(ZMA); Santubong, Sarawak, Borneo (both USNM); Labuan,
Borneo (NMW, RNHL); Sipitang, North Borneo; Po Bui
Island, Sandakan, North Borneo (both USNM).
[05-339]
466 Littorinopsis
?Littorina miodelicatula (Oyama, 1950)
(Pl. 358, figs. 1-3)
Range—Tertiary of Japan.
Remarks—The_ species was originally de-
scribed as resembling some of the Trochidae but
Oyama preferred an assignment in Littorinidae
near L. scabra because of residual color pattern in
the fossil and a shape somewhat like that of L.
carinifera. It is included here somewhat doubt-
fully as the affinity with Archaeogastropoda
seems most likely.
Synonymy—
1950 Littorinopsis (Littorinopsis) miodelicatula Oyama, Geo-
logical Survey of Japan, Report no. 132, p. 8, pl. 1,
figs 2, 3. (Ueno, Japan; Tertiary Kakebata formation)
holotype in collection of Geological Survey of Japan
[?]: 8 x 6.7 mm.
Littorina incisa Yokoyama, 1927
(Pl. 358 figs. 4, 5)
Range—Pliocene of Japan.
Remarks—Littorina incisa is a relatively small
but well-preserved fossil. The excellent illustra-
tion given by Yokoyama and reproduced here
indicates that this species is probably most close-
ly related to L. melanostoma and is quite unlike
any other Recent species because of its attenuate
shape.
Synonymy—
1927 Littorina incisa Yokoyama, Journal of the Faculty of
Science, Imperial University of Tokyo, section II,
Geology, Mineralogy, Geography, Seismology, vol. 2,
part 4, p. 175, pl. 47, fig 8 (Pliocene, Nagaya, Kaga,
Japan); holotype in Geological Institute, Imperial
University of Tokyo: 5 x 2.5 mm.
Joseph Rosewater
Littorinidae
Plate 358. Figs. 1-3. Littorinopsis miodelicatula Oyama, from
Tertiary, Ueno, Japan (Holotype, from original figures in
Report no. 132, Geological Survey of Japan, pl. 1, figs.
3a-c; 8 x 6.7 mm.).
Figs. 4,5. Littorina incisa Yokoyama, from Pliocene,
Nagaya, Kaga, Japan (Holotype from original figures; 5 x
2.5 mm.).
[05-340]
November 30, 1970
Subgenus Austrolittorina new subgenus
Rosewater
Type-species: Littorina unifasciata unifasciata Gray, 1826
The members of this subgenus are character-
ized by a predominantly southern ocean and
tropical distribution and by the possession of
shell and anatomical features generally similar to
those of its type-species, L. unifasciata unifas-
ciata: a conico-turbinate shell, flattened columel-
la and crescent-shaped area on the adjacent part
of the shell; verge with a basal enlargement
incorporating a single penial gland containing an
accessory flagellum.
Littorina unifasciata Gray, 1826
The Littorina unifasciata species complex is
represented in the Indo-Pacific faunal region by
three entities, the nominate subspecies, unifas-
ciata in Australia, and the subspecies antipoda
in New Zealand and fernandezensis in the Juan
Fernandex Islands. All closely resemble each
other except for size or proportional differences
which are the result, probably, of isolation. All
members of the subgenus Austrolittorina show a
preference for the southern ocean south of the
equator.
Littorina unifasciata
subspecies unifasciata Gray, 1826
(Pls. 325, 326, 359-361)
Range—Southern coast of Australia, predomi-
nantly south of the Tropic of Capricorn, from
Carnarvon in the west to Queensland; Lord Howe
Island; Tasmania.
Remarks—The nominate subspecies of the L.
unifasciata group is distinctly a native of the
Australian continent occurring extraterritorially
only in Tasmania and on Lord Howe Island. The
plot of its distribution, if smoothed, would resem-
ble an inverse normal curve (pl. 362); it is re-
stricted almost exclusively to the more southern
coasts south of the Tropic of Capricorn. This
species has been confused in the literature with
L. mauritiana Lamarck, although the two are
quite distinct morphologically and their ge-
ographic ranges are separate (see mauritiana).
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 467
Both Reeve (1858) and Tryon (1887) considered
unifasciata to be a synonym of mauritiana. Al-
though both may have the single dark band
around the whorls upon which the name “unifa-
sciata” is based, mauritiana is often larger and
more elongate, it bears subtle darker patterning
and lacks the columellar crescent and _ penial
gland of unifasciata.
Habitat—Lives on shore rocks in the spray
zone, above high water.
Description—Shell reaching 20.9 mm. (about .8
inch) in length, conical to subturbinate in shape,
average obesity about .62 (45 specimens range
from .55 - .70); older individuals moderately thick
in structure, imperforate, usually developing a
flattened, crescent-shaped area adjacent to the
columellar callus; sculptured with spiral striae
occasionally reinforced with fine brown color
lines, spiral, white microscopical textural threads,
and irregular, oblique, axial lines of growth. Ex-
ternal ground color grayish white to bluish gray,
usually the former with a rather diffuse blue-gray
band encircling the body whorl and anterior
portion of spire whorls; apex light brown. Aper-
ture medium to dark-brown with a prominent
white band inside near junction of outer lip and
columella. Base somewhat flattened, separated
from upper part of body whorl by a low but
distinct keel at periphery. Whorls 5-7, rather
straight-sided. Spire less than half the length of
shell, convex, produced at an angle of about 60°.
Aperture oval to subquadrate; outer lip moder-
ately thick, having its origin high on body whorl
above keel so that keel enters aperture; outer lip
striated internally at edge, reflecting external
sculpture; inner lip weakly concave. Columella
dark brown to white, shallowly excavated, some-
what rimmed medially, and with a flattened
brown to white crescent-shaped area distally on
the adjacent base. Suture not deeply impressed.
Sculpture consisting of from 6-9 spiral striae on
spire whorls, persisting onto surface of body
whorl where as many as 12 may be present above
the keel; becoming indistinct below keel. Entire
surface covered with microscopic, closely spaced
rather straight spiral white threads partially im-
bedded in shell substance and seen through the
surface; not detectable in worn specimens. Axial
sculpture consisting of rather irregular oblique
lines of growth. Operculum corneous, moderately
thick, paucispiral. Periostracum not evident in
specimens examined. Nuclear whorls decollate in
most mature specimens; when present, light
brown in color, smooth and about 3 in number;
[05-351]
468 Austrolittorina
first post-nuclear whorl similarly colored, but
sculptured with spiral striae. Radula littorinid
(2-1-1-1-2) central tooth somewhat narrow.
Animal darkly pigmented on surfaces of ten-
tacles, snout and foot. Verge moderately short and
thick, yellowish white in color; having a basal
Joseph Rosewater
Littorinidae
flap with a single penial gland containing an
internal hyaline accessory flagellum. Nothing is
known concerning the reproduction and develop-
ment of this species, although probably it is
oviparous and spawns pelagic capsules.
Plate 359. Subspecies of Littorina (Austrolittorina) unifasciata
(Gray).
Figs. 1,2. L. unifasciata unifasciata from Port Jackson,
New South Wales, Australia (USNM 89472; 20.9 x 12
mm.; note flattened crescent sloped area adjacent to colu-
mellar callous).
Figs. 3,4. L. unifasciata unifasciata from Red Bluff, Kalbar-
ri, Western Australia (USNM 691673; 12.7 x 7.6 mm.).
Fig. 5. L. unifasciata unifasciata from same locality as
Figs. 3,4; detail of shell surface to show white microscop-
ical textural threads (enlargement of shell area measuring
about 1.3 x .8 mm.; young specimen, length 4.9 x 3.2
mim.).
Figs. 6-8. L. unifasciata antipodum; fig. 6, from Island
Bay, Cook Strait, New Zealand (USNM 671202, 8.7 x 4.8
mm.); figs. 7,8, from Taipa, Doubtless Bay, North Island,
New Zealand (USNM 601974; 7.4 x 4.7 mm.).
Figs. 9-12. L. unifasciata fernandezensis from eastern shore
Cumberland Bay, Isla Mas a Tierra figs. 9, 10, Paratype
(USNM 679256, 11.8 x 7.2 mm.); figs. 11, 12, Holotype
(USNM 368900, 13.4 x 7.8 mm.),
[05-352]
November 30, 1970
Measurements (mm.)—
length width no. whorls locality
20.9 11.9 5+ Port Jackson,
New South Wales
19.9 12.9 5+ Port Jackson,
New South Wales
19.3 11.3 5 Port Jackson,
New South Wales
18.2 11.4 6 Kalbarri, Western
Australia
17.9 11.6 4+ Kalbarri, Western
Australia
16.3 9.5 5+ Robe, S. E. Australia
16.0 9.6 5 Port Denison,
Western Australia
15.3 9.3 5+ Port Fairy, Victoria
14.3 9.1 5 Mouth of Murchison
River, Western
Australia
13.0 7.9 4 near Sydney,
New South Wales
1:29 8.1 4 Mouth of Murchison
River, Western
Australia
10.9 6.5 4+ Queenscliff, Victoria
10.7 Wee 5 Wollongong, N.S.W.
9.4 6.1 4 Coogie Beach,
Sydney, N.S.W.
Heo 4.7 5 Coogie Beach,
Sydney, N.S.W.
6.4 3.8 3+ Coogie Beach,
Sydney, N.S.W.
5.0 Sill 3+ Pearl Beach, N.S.W.
4.9 3.4 3+ between Port Phillip
Heads and Cape
Otway, Victoria
Synonymy—
1826 Littorina unifasciata Gray, in P. P. King, Narrative of a
survey of the coasis of Australia, vol. 2, Appendix B,
p. 483 (Australia [King George Sound, Western Aus-
tralia, here restricted ]); lectotype B.M.(N.H.) 1968373:
16.4 x 10.6 mm.
1833 Littorina diemenensis Quoy and Gaimard, Voyage de
decouvertes de ]’Astrolabe, vol. 2, part 2, p. 479, pl.
33, figs. 8-11 (rocks of the littoral zone of South
Australia, Tasmania and also New Zealand [here
restricted to Tasmania]); lectotype in Mus. d Hist.
Nat., Paris: 10.2 x 5.9 mm.
1843 Litorina acuta Menke, Molluscorum Novae Hollandiae
Specimen, p. 9 (Western Australia; type lost); 1844,
Zeitschrift fiir Malakozoologie, Jahr, 1844. p. 57.
1847 Litorina mauritiana crassior Philippi, Abbildungen und
Beschreibungen Conchylien, vol. 2, p. 165, Litorina
pl. 3, fig 17[a] (Australia [here restricted]).
1850 Littorina diemensis Gray, Figures of Molluscous Ani-
mals, vol. 4, p. 78 [error for L. diemenensis Quoy and
Gaimard, 1833].
1858 Littorina laevis ‘Philippi’ Reeve, Conchologia Iconica,
vol. 10, Littorina, pl. 17, fig 95 (locality not given);
not L. laevis Philippi = L. mauritiana Lamarck.
1885 Littorina diemenensis pseudolaevis Nevill, Hand-List of
Mollusca in the Indian Museum, Calcutta, part 2, p.
141, (New South Wales); new name for L. laevis
Reeve, not L. laevis Philippi.
Types—The lectotype of L. unifasciata Gray is
in the BM(NH), 1968373. The type locality men-
tioned by Gray, Australia, is here restricted to
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 469
Plate 360. Figs. 1,2. Lectotype of Littorina unifasciata Gray
from Australia [King George Sound, Western Australia] (B.
M.(NH) 1968373, 16.4 x 10.6 mm.).
Figs. 3,4. Lectotype of Littorina diemenensis Quoy and
Gaimard [= L. unifasciata] [from Tasmania] (MHNP,
10.2 x 5.9 mm.).
Fig. 5. Lectotype figure of Litorina antipodum Philippi,
from Abbildungen und Beschreibungen Conchylien, vol.
2, p. 195, pl. 4, fig. 2 (New Zealand, about 7.8 x 4.5 mm.).
King George Sound, Western Australia, one of the
localities visited during King’s survey. The lecto-
type of L. diemenensis Quoy and Gaimard is in
the Paris Museum. It is suspected that the illus-
tration is a composite since there were no speci-
mens in the type lot which fully resembled it. A
lectotype was chosen which most closely approxi-
mated the illustration, although smaller than the
original measurements (10.2 x 5.9 mm. versus
approximately 11.3 x 6.8 mm. (5x 3 lignes)). The
holotype of Menke’s L. acuta may be presumed
lost; however the author himself (1844, see Syn-
onymy) synonymized his species with unifasc-
iata an action which is accepted here as proof of
the identity of the species in view of the absence of
a type specimen. The type of L. crassior Philippi
may be in the Berlin Museum; the type of L.
pseudolaevis Nevill may be in the Indian Muse-
um, Calcutta.
Records—AUSTRALIA: QUEENSLAND: Point Lookout,
Stradbroke Island, Moreton Bay (WAM, AMS); Freshwater;
Noosa Heads; Caloundra; Coolangatta (all AMS). NEW
SOUTH WALES: Byron Bay (MCZ, AMS); Yamba (MCZ);
Merewether Beach, Newcastle (WAM); Pearl Beach: The
[05-353]
470 Austrolittorina
AI
AP YT
; 5mm
Plate 361. Fig. A. Radula of Littorina unifasciata unifasciata
from The Entrance, New South Wales, Australia, (USNM
631284; upper scale is .05 mm.)
Fig. B. Radula of L. unifasciata fernandezensis from Mas
Afuera Id., Juan Fernandez Islands (middle scale is .1
mm.).
Fig. C. Penis of L. unifasciata unifasciata from Kalbarri,
Western Australia, USNM 691673 (lower scale is 5 mm.).
Entrance (both USNM); Putty Beach (AMS): S. shore, Broken
Bay, nr. entrance (ANSP); btwn. Newport and Palm Beach
(ANSP, USNM; Middle Harbor (AMS); Long Reef, N. of
Manly, nr. Sydney (AMS, ANSP); Sydney (MCZ): Port Jack-
son (USNM, MCZ, AMS, ANSP); La Perouse, Botany Bay
(MCZ, ANSP, WAM); Kurnell (AMS); Wollongong (AMS,
USNM); Port Kembla (USNM); Merimbula; Two Fold Bay
(both MCZ, AMS), VICTORIA: Mallacoota (AMS); Flinders
(USNM); Sandringham, Port Philip (ANSP, USNM):; Port
Philip (ANSP, AMS, MCZ); btwn. Port Philip Heads and
Cape Otway (USNM); Barwon Heads (MCZ); Port Fairy
(ANSP, USNM, AMS, NMW); Bridgewater Bay (MCZ); Port-
land (MCZ); TASMANIA: Fischer Island, Bass Strait (AMS);
Islets S. of Flinders Island, Bass Strait (WAM); King Island;
Bicheno; Northwest Long Bay, Port Arthur; Simpson’s Bay,
D’Entrecasteaux Channel (all AMS); Eagle Hawk Neck; Ho-
bart (both MCZ); Taroona, Derwent River; Grant’s Head,
Sloop Rocks (both AMS); Low Head (MCZ). SOUTH AUS-
TRALIA: Boatswain Point (Cape Thomas) nr. Robe (ANSP,
USNM, MCZ); Robe (NMW); Encounter Bay; Port Willunga
(both MCZ); Adelaide (USNM); Giles Point, Yorke Peninsula
(USNM, MCZ, NMW); Wool Bay (NMW); South Neptune
Island, Spencer Gulf (MCZ, AMS). WESTERN AUSTRALIA:
Mondrain Island, Recherche Archipelago (WAM); Esperance
(AMS); mouth of Pallinup R. Estuary (WAM); Frenchman’s
Bay, nr. Albany (AMS); Oyster Harbor, Nr. Albany, N. of
Bayonet Head; Middleton Bay, nr. Albany; Windy Harbor, Nr.
Northcliffe (all WAM); Cape Leeuwin, (AMS); Margaret River
(WAM); Wallcliffe, nr. Margaret River (MCZ); Yallingup,
Cape Naturaliste; Bunker Bay, Cape Naturaliste (both
USNM); Geographe Bay; Bunbury (both AMS); Cottesloe
(WAM, AMS): Point Peron, nr. Perth, on boulders; Rottnest
Island (both MCZ, AMS); Perth (AMS); Little Anchorage,
Leeman; Snag Island, Leeman; Beagle Islands (all WAM);
Port Denison (USNM); Greenough River mouth; Harrocks
Beach nr. Northampton (both WAM): Red Bluff, Kalbarri, nr.
mouth of the Murchison River (WAM, USNM); Eagle Bluff,
Freycinet Estuary (WAM); Point Gregory, N.W. Corner Peron
Peninsula (WAM); Dirk Hartog Island, (last three all Shark
Bay) (AMS); below Quodd’s Light, N. of Carnarvon (WAM),
LORD HOWE ISLAND (ANSP, AMS).
Joseph Rosewater Littorinidae
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Poy :
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Norfolk
Plate 362. Geographical distribution of Littorina (Austrolit-
torina) unifasciata unifasciata (Gray), in Australia, and of the
subspecies, L. unifasciata antipodum (Philippi), in New Zea-
land.
Littorina unifasciata
subspecies antipodum (Philippi, 1847)
(Pl. 359, figs. 6-8, pl. 360)
Range—Islands of New Zealand.
Remarks—The New Zealand species best
known as Littorina oliveri (Finlay) is without a
doubt L. antipodum Philippi, here considered a
subspecies of L. unifasciata. Finlay’s (1930) com-
parison of antipodum with the typical Australian
unifasciata best summarizes the differences:
“The New Zealand species differs in its smaller
shell, higher and more slender spire, with almost
straight instead of distinctly convex whorls, no-
tably smaller aperture, and darker more promi-
nent and better defined blue band .. .” It may be
theorized that elements of unifasciata were car-
ried to New Zealand in the past and that the
subspecific differences evolved in response to the
new environmental conditions during isolation.
Habitat—On rocks at and above high water
mark.
Description—Shell reaching 12.3 mm. (about .5
inch) in length; in general appearance looking
much like a diminutive L. unifasciata unifas-
ciata, elongate to rather short-conical in shape;
average obesity about .59 (21 specimens range
from .52-.64). A flattened, crescent-shaped area
adjacent to columella callus usually present, but
often reduced in size or may be lacking. Sculpture
similar to L. unifasciata unifasciata. Color band
encircling body whorl and anterior portion of
[05-354]
November 30, 1970
Plate 363. Geographical distribution of Littorina (Austrolit-
torina) unifasciata fernandezensis Rosewater in the Juan Fer-
nandez Islands, southeastern Pacific.
spire whorls very dark bluish-gray, shell often
tinted brown above and below the band. Whorls
4-6, apex usually severely eroded. Radula as in L.
unifasciata, the central tooth rather narrow.
Animal darkly pigmented on surfaces of ten-
tacles, snout and foot. Verge similar to L. cincta
and L. unifasciata unifasciata: rather short and
thick and having a basal mitten-shaped flap
which bears a penial gland with an internal
hyaline accessory flagellum. Sexually mature
males very small. Nothing known concerning
reproduction and development, although is prob-
ably oviparous and spawns pelagic capsules.
Measurements (mm.)—
length width no. whorls locality
are 6.9 6+ Wellington,
New Zealand
123 6.4 54 Wellington,
New Zealand
10.3 5.6 6+ Port Waikato,
New Zealand
10,2 6.0 6 Wellington,
New Zealand
9.7 6.0 44 Christ Church,
New Zealand
9.5 6.1 44 Doubtless Bay, North
Island, New
Zealand
8.8 5.0 44 Island Bay, Cook Str.
New Zealand
8.5 ee 5 Napier, New Zealand
7.9 5.0 6 “New Zealand”
5.4 3.2, 5 Auckland,
New Zealand
Synonymy—
1847 Litorina antipodum Philippi, Abbildungen und Besch-
reibungen Conchylien, vol. 2, p. 195, Litorina pl. 4,
fig 2 (New Zealand)
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 471
1930 Melarhaphe oliveri Finlay, Transactions of the New
Zealand Institute, vol. 61, p. 224, (Hampden, East
Coast of South Island, New Zealand; Holotype in
Finlay Collection, Auckland Museum); refers to
Suter, 1913, p. 188, and to C. E. R. Bucknill, 1924,
“Sea Shells of New Zealand,” p. 37, pl. 7, no. 2); not
Melarhaphe zelandiae Finlay, 1926, which = L.
cincta Quoy and Gaimard.
Types—The type of L. antipodum Philippi may
be in the Berlin Museum. The lectotype figure is
that of Philippi, Abbildungen und_ Beschrei-
bungen Conchylien, vol. 2, Litorina, pl. 4, fig 2
(see pl. 360 fig. 5). The holotype of M. oliveri
Finlay is in the Finlay Collection, Auckland
Museum. It should be pointed out here that
Finlay (1926) by error described Melarhaphe ze-
landiae intending the name for the New Zealand
relative of L. unifasciata. However, the type of
zelandiae proved to be identical with L. cincta,
another New Zealand species and it was for this
reason that Finlay (1930) thought it necessary to
describe M. oliveri not realizing that Philippi had
already described the species over 80 years be-
fore.
Records—NEW ZEALAND: NORTH ISLAND: Tauranga
(BPBM); Napier; Plimmerton, Wellington; Titahi Beach, in
shell sand (all USNM); Island Bay, Cook Strait (AMS,
USNM); Worser Bay, Wellington Harbour (MCZ, USNM);
New Plymouth (AMS); Taipa, Doubtless Bay (USNM); Rus-
sell, Bay of Islands (MCZ); Hen Island; N.W. point on
Motutapu Island (both ANSP); Auckland (AMS, ANSP); W.
side Wade River, about 15 mi. N.W. of Auckland (USNM,
ANSP); Rangitoto (AMS, ANSP); S. coast, Rangitoto Island
(USNM); Manukau Harbor; Port Waikato (both ANSP,
USNM). SOUTH ISLAND: Picton, Lyttelton (AMS); Taylor's
Mistake, Christchurch (USNM); Porto Bello; Dunedin; Gray-
mouth (all AMS); Stewart Island (ANSP); Chatham Islands
(Suter, 1913).
Littorina unifasciata
new subspecies fernandezensis Rosewater
(Pls. 359, 361)
Range—Juan Fernandez Islands, southeastern
Pacific.
Remarks—Odhner (1922) reported Littorina
mauritiana Lamarck from Mas Afuera, Juan Fer-
nandez Islands, and referred to its distribution as
extending from Mauritius to New Zealand. In the
present study it has been found that L. mauri-
tiana is a species restricted to the western Indian
Ocean, and that L. unifasciata unifasciata and its
subspecies L. u. antipodum, inhabiting Australia
and New Zealand respectively, are quite distinct.
The third subspecies of the L. unifasciata group,
named here, is located geographically quite far
from its nearest relative. According to Odhner’s
[05-355 ]
472 Austrolittorina
Joseph Rosewater
Littorinidae
(ibid.) analysis the faunal affinities of Juan Fer-
nandez show a closer relationship to South Amer-
ica than to the western Pacific. Further, he found
high endemism in the mollusks, indicating long
isolation. Nevertheless the Littorina from Juan
Fernandez is clearly related to L. unifasciata, but
with sufficient differences to suggest that at least
subspeciation has occurred.
The answer to the question of how L. unifas-
ciata and its subspecies became distributed in
such widely separated localities will probably
never be answered satisfactorily. Although I
know of no fossil record, the history of the
ancestors of this group probably dates well back
into geologic time when land masses and ocean
currents are known to have been quite different. It
is possible that the group evidences a relict and
disjunct distribution from the time when the
Antarctic continent was both larger and warmer
than it is today. The subspecies living in southern
Australia, New Zealand and Juan Fernandez may
have evolved from a once more widely dis-
tributed species which migrated northward in
response to the cooling off of the southern con-
tinent. It is also possible that they represent a
modern distribution brought about by the pre-
dominately western current of the southern
ocean. (See discussion of a somewhat similar
circumpolar distribution by Abbott, 1968, pp.
183-188).
Littorina unifasciata fernandezensis is very
similar to the nominate subspecies, differing in
being on the average slightly more slender, al-
though it is not so slender as L. u. antipodum (see
average obesities in Descriptions of L. unifas-
ciata and antipodum). There is a superficial simi-
larity in appearance between L. fernandezensis
and L. paytensis Philippi of western South Amer-
ica, but the former lacks the brown color mark-
ings, two white bands in the aperture and single
strong subsutural stria which are all present in
paytensis. The two also are members of different
subgenera of Littorina!
Habitat—Shore rocks.
Description—Shell reaching nearly 18 mm.
(about .7 inch) in length, conical to subturbinate
in shape average obesity about .60 (52 specimens
ranged from .55 - .64); appearance very similar to
L. unifasciata unifasciata, but more turbinate in
shape and less conical; columella not as strongly
developed; adjacent crescent-shaped, flattened
area persists and may be well developed in some
specimens. Spiral striae often strongly impressed,
16-22 striae on body whorl above periphery (14-
16 in L. unifasciata unifasciata); striae not rein-
forced by brown color lines; spiral sculpture
sometimes apparent inside edge of outer lip as a
series of fine teeth. Color of aperture varying from
brown to violet; color band on body whorl often
narrower and darker than in unifasciata. Animal
characters similar to L. unifasciata unifasciata
Measurements (mm.)—
length width no. whorls locality
17.6 11.3 34 Santa Clara Id.
Islas Juan Fernandez
16.3 10.2 5+ Cumberland Bay,
Isla Mas a Tierra
(paratype)
15.7 9.0 5+ Cumberland Bay
15.3 9.9 4+ Santa Clara Id.
14.4 8.3 6 Cumberland Bay,
Isla Mas a Tierra
(paratype)
13.4 7.8 5 Cumberland Bay,
(holotype)
13.4 7.9 5+ Isla Mas Afuera
13.2 8.2 44 Santa Clara Id.
13.0 8.1 5+ Isla Mas Afuera
12.8 8.0 4+ Isla Mas Afuera
12.4 7.3 6 Isla Mas Afuera
11.9 7.4 5 Cumberland Bay,
Isla Mas a Tierra
(paratype)
11.1 6.8 44 Isla Mas Afuera
10.5 6.6 44 Cumberland Bay,
Isla Mas a Tierra
10.1 6.5 54 Isla Mas Afuera
9.8 Dal. 44 Isla Mas Afuera
8.6 5.4 5+ Isla Mas Afuera
7.0 4.3 54 Cumberland Bay,
Isla Mas a Tierra
(paratype)
6.2 oer 5+ Cumberland Bay
5.2 3.0 44 Isla Mas Afuera
Types—The holotype was collected by Dr.
Waldo L. Schmitt from the littoral zone, east
shore of Cumberland Bay, Isla Mas a Tierra, Juan
Fernandez Islands, in 1926 (USNM_ 368900);
there are 25 paratypes (USNM 679256).
Synonymy—
1885 Littorina penitaria Wood, in G. Nevill, Hand List of
Mollusca in the Indian Museum, Calcutta, part 2, p.
142 (San Juan Fernandez [sic]); [Nomen nudum].
1922 Littorina mauritiana Lamarck, in N. H. Odhner, The
Natural History of Juan Fernandez and Easter Island,
edited by Dr. Carl Skottsberg, vol. 3, part 2, p. 223
(Isla Mas Afuera; not L. mauritiana (Lamarck,
1822).
Records—ISLAS JUAN FERNANDEZ: Isla Mas Afuera
(Odhner, 1922); East side of Isla Mas Afuera (Eltanin Cruise
21, Ann Cohen, Collector; USNM); Holotype and 25 para-
types, east shore Cumberland Bay, Isla Mas a Tierra; North
Bay, Isla Santa Clara (both collected by W. L. Schmitt in 1926;
USNM). Paratypes collected by William Dodd in Del. Mus.
Nat. Hist. no. 39221.
[05-356 ]
November 30, 1970
Littorina cincta Quoy and Gaimard, 1833
(Pls. 364, 365)
Range—North, South and Stewart Islands, The
Snares and Chatham Islands, New Zealand.
Remarks—There is little reason for confusing
the two larger species of Littorina which inhabit
the shores of New Zealand. The closely brown-
banded L. cincta offers a rather striking contrast
Plate 364. Figs. 1-6, Littorina (Austrolittorina) cincta (Quoy
and Gaimard).
Figs. 1,2. Lectotype of Littorina cincta, from New Zealand
(MHNP; 12.1 x 6.8 mm.).
Figs. 3,4. Holotype of Littorina luctuosa Reeve (BM(NH)
1968315, 13.7 x 7.6 mm.).
Figs. 5,6. Holotype of Melarhaphe zelandiae Finlay (from
Transactions of the New Zealand Institute, 1926, vol. 57,
pl. 18, figs. 18, 19; 17 x 10.5 mm.).
Figs. 7-10. Littorina (Austrolittorina) punctata (Gmelin).
Figs. 7,8. Littorina punctata, from 20 km. south of Luanda,
Angola, West Africa (USNM 679288; 9.3 x 6 mm.).
Figs. 9,10. Holotype of Turbo punctatus Gmelin, Senegal
(from Journal de Conchyliologie, 1942, vol. 85, pl. 10,
figs. 2a, 2b; MHNP, 17 x 11 mm.).
Figs. 11-16. Littorina (Austrolittorina) africana (Philippi).
Figs. 11,12. Lectotype of Litorina africana, from Cape of
Good Hope, South Africa (Stuttgart Museum (NH): MT
106, 9.8 x 7.3 mm.; photo is from Janus, 1961, Stuttgarter
Beitrage zur Naturkunde, no. 70, pl. 3, figs. 1,2).
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 473
to the lighter, grayish blue L. unifasciata anti-
podum which has a single dark and often diffuse,
revolving color band on each whorl. Although
there is a superficial resemblance in color pattern
between L. cincta and L. pintado, details of
anatomy and of shell morphology confirm their
distinctness.
Habitat—On rocks at and above high tide line.
Description—Shell reaching 20 mm. (Suter,
1913; about 0.8 inch) in length, conic-turbinate in
shape, average obesity about .60 (23 specimens
range from .52-.67); older individuals only
moderately thick in structure, imperforate, usual-
ly developing an often rather narrow flattened,
crescent-shaped area adjacent to columellar cal-
lus; sculptured with often faint spiral striae rein-
forced by white spiral lines to a varying degree;
fine to rather coarse axial lines of growth present.
External color pattern consisting of the narrow,
white spiral bands with wider medium to dark
brown spiral bands interspersed. Aperture usual-
ly dark-brown, the narrow white lines showing
through inside outer lip, with a broad white band
near junction of outer lip and columella. Base
hardly flattened, separated from upper part of
body whorl by a rather weak keel at periphery.
Whorls 5-7, moderately rounded. Spire less than
half the length of shell, convex, produced at an
angle of about 55°. Aperture oval; outer lip rather
thin, inner lip weakly concave. Columella brown
to white, shallowly excavated, somewhat rimmed
medially and with a flattened crescent-shaped
area distally on the adjacent base. Suture im-
pressed. Operculum corneus, paucispiral. Perios-
tracum not evident in specimens examined.
Nuclear whorls smooth and colorless; first 2-3
post-nuclear whorls dark-brown, unsculptured,
Figs. 13,14. Littorina africana, from Tiger Rocks, Isipingo,
Natal, South Africa (USNM 637358, 9.4 x 6.5 mm.).
Figs. 15,16. Lectotype of Litorina decollata Philippi, from
Natal, South Africa (Stuttgart Museum (NH): MT 107, 5.8
x 4.3 mm.; photo from Janus 1961, Stuttgarter Beitrage
zur Naturkunde, No. 70, pl. 3, figs. 3,4).
Figs. 17-23. Littorina (Austrolittorina) knysnaensis (Philip-
pi).
Figs. 17,18. Lectotype of Litorina knysnaensis from Cape
of Good Hope, South Africa, near Knysna River (Stuttgart
Museum (NH): MT 108, 9.3 x 6.0 mm.; photo from
Stuttgarter Beitrage zur Naturkunde, no. 70, pl. 3, figs.
5,6).
Figs. 19,20. Lectotype of Littorina picea Reeve [from South
Africa] (BM(NH) 1968320, 6.9 x 4.5 mm.).
Figs. 21-23. Littorina knysnaensis from “South Africa’; in
fig. 22, basal view, and in fig. 23, note spotted color
pattern particularly characteristic of base of shell in this
species (USNM 633262, 11.1 x 6.9 mm.).
[05-357 ]
474 Austrolittorina
shining. Radula littorinid (2-1-1-1-2); similar
to L. unifasciata central tooth somewhat narrow.
Animal darkly pigmented on upper surfaces of
tentacles, snout and foot. Verge short and thick,
yellowish white in color; having a large basal flap
which bears a single penial gland containing an
internal hyaline accessory flagellum. Reproduc-
tion unknown, probably oviparous with pelagic
capsule.
Measurements (mm.)
length width no. whorls locality
19.2 11.0 6+ Wellington Harbour,
New Zealand
18.8 10.6 6 Caroline Bay, Timaru,
South Island
18.2 9.8 6+ Manukau Harbour
17.1 10.3 4+ Solander Id.,
Foveaux Strait
16.1 9.3 5+ Auckland
15.4 8.0 6+ Stewart Island
14.4 8.1 5+ Stewart Island
13.5 9.1 4+ Caroline Bay, Timaru,
South Island
13.3 8.4 4+ Bay of Islands
11.8 7.1 5 Stewart Island
es 7.1 4+ Worser Bay
9.2 6.0 4+ Stewart Island
8.8 5.4 4+ Auckland
Synonymy—
1833 Littorina cincta Quoy and Gaimard, Voyage De L’Astro-
labe vol. 2, part 2, p. 481, pl. 33, figs 20-21 (New
Zealand); lectotype in Museum d Histoire Naturelle,
Paris: 12.1 x 6.8 mm; not L. cincta Gould, 1847,
Proceedings of the Boston Society of Natural History,
vol, 2, p. 252 (from Puget Sound) [=L. sitchana
Philippi].
1857 Littorina luctuosa Reeve, Conchologia Iconica, Vol. 10,
Littorina, pl. 13, fig. 65 (New Zealand); Holotype
BM(NH) 1968315: 13.7 x 7.6 mm.
1926 Melarhaphe zelandiae Finlay, Transactions of the New
Zealand Institute, vol. 57, p. 375, pl. 18, figs. 18, 19
[description and figs. = L. cincta] (Dunedin Har-
bour; type in Finlay Collection, Auckland Museum,
17 x 10.5 mm.); erroneously proposed as a new
species name for Littorina mauritiana ‘Lamarck’
Suter, 1913, p. 188, which = L. unifasciata anti-
podum Philippi (see Finlay, 1930, Transactions of
the New Zealand Institute, vol. 61, p. 224).
Types—A lectotype of Littorina cincta Quoy
and Gaimard is here designated from among three
syntypes in the Museum National d’Histoire Na-
turelle, Paris. (see pl. 364 figs. 1, 2). It is suspected
that the figure in L’Astrolabe Atlas is a composite
drawing as none of the syntypes match it exactly
and it is smaller than the measurement accom-
panying the description: 12.1 x 6.8 mm. vs 6 x 4
lines [= about 13.5 x 9.0 mm.].
The holotype of Littorina luctuosa Reeve is in
the British Museum (NH) 1968315 (pl. 364, figs.
3, 4). The type of Melarhaphe zelandiae Finlay is
in the Auckland Museum (pl. 364, figs. 5, 6).
Joseph Rosewater
Littorinidae
Records—NEW ZEALAND: NORTH ISLAND: Plim-
merton, Wellington (USNM); Worser Bay (USNM, MCZ);
Oriental Bay, both Wellington Harbor (USNM); Wanganui
(MCZ); New Plymouth (AMS); Doubtless Bay; Russell (both
MCZ); Long Beach, both Bay of Islands (USNM); Hen Island,
off E. coast Auckland Province (MCZ); Auckland (USNM,
ANSP, MCZ); Muriwai Beach (MCZ). SOUTH ISLAND:
Lyttelton; Akaroa Banks Peninsula (both AMS); Picton;
Kaikoura; Timaru (all MCZ); Caroline Bay, Timaru (USNM);
Kartiki, Otago (ANSP); Purakanui, N. Otago (MCZ); Hatch-
ery, Portobello; Pipikariti, Dunedin; Wangaloa; Riverton (all
AMS); Wangaloa; Riverton (all AMS); Point Elizabeth (MCZ);
Greymouth (AMS, MCZ). FOVEAUX STRAIT: Solander Is-
land (USNM). STEWART ISLAND (USNM, ANSP). CHAT-
HAM ISLANDS. THE SNARES (both Suter, 1913).
Littorina punctata (Gmelin, 1791)
(Pls. 364, 365)
Range—The Mediterranean, West and South
Africa.
Remarks—Although not truly a member of
Indo-Pacific Littorinidae, L. punctata is a tropical
species which appears to belong in the subgenus
Austrolittorina and evidences many characteris-
tics similar to L. unifasciata. Its shell is often
subturbinate or conical, there is a tendency to
Plate 365. Fig. A. Penis of Littorina cincta Quoy and Gaimard,
from Plimmerton, Wellington, New Zealand (USNM 671204).
Fig. B. Penis of Littorina punctata (Gmelin) from Goree,
Daker, Senegal (ZMC).
Fig. C. Penis of Littorina africana (Philippi) from 11 miles
south of Port Shepstone, Natal, South Africa (ANSP
216678); shading indicates pigmentation noted in this and
some other specimens.
Fig. D. Penis of Littorina knysnaensis (Philippi) from N.E.
False Bay, Cape Province, South Africa (ANSP 216643);
dark area indicates pigmentation noted in some speci-
mens. All drawn to same scale.
[05-358]
November 30, 1970
develop a semilunar depression adjacent to the
columella, a white band revolving into the aper-
ture, and the verge has a flap with a single hyaline
spur. The range of L. punctata is quite extensive:
from the Mediterranean to South Africa, via West
Africa. The species may be recognized readily by
the overall pattern of white spots on a dark
background from which its name is derived.
It is included here to avoid confusion with
Indo-Pacific species whose ranges extend into or
are limited to South African waters.
Habitat—Shore rocks, in the spray zone.
Description—Shell may exceed 18 mm. (about
0.7 inch) in length, conical to subturbinate in
shape; average obesity about .64 (26 specimens
range from .61-.68) moderately thick in struc-
ture, imperforate, usually developing a narrow,
flattened, crescent-shaped area adjacent to colu-
mella callous; sculptured with only moderately
well-impressed spiral striae and irregular axial
growth lines. External ground color light to dark
brown, with an overall pattern of white spots
varying in shape from small, compact rhomboidal
to elongate. Aperture medium to dark brown with
a prominent white band near junction of outer lip
and columella. Base somewhat flattened, sepa-
rated from upper part of body whorl by a low but
distinct keel at periphery. Whorls 5-7, rather
straightsided. Spire less than half the length of
shell, convex, produced at an angle of about 60°.
Aperture oval; outer lip moderately thick, having
its origin high on body whorl so that keel enters
aperture; inner lip weakly concave. Columella
tannish white, shallowly excavated, stout appear-
am ™ a] |
> he: %
ae f OL. Ritts. | = |
aoe ee
| NEW SQhoR +, TONGA p
Fel eee : “LU CALeDONia_ Os is. _ COOK] -
. Norfo/k
[CINCTA]
140° 160° 180° 160°
Plate 366. Geographical distribution of Littorina (Austrolit-
torina) cincta (Quoy and Gaimard), in New Zealand.
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 475
ing and with a narrow crescent-shaped area on
adjacent base. Suture not deeply impressed. Spire
sculpture often obscured by wear; 15-17 spiral
striae on body whorl above keel, becoming ob-
scure below on base. Operculum corneous, pau-
cispiral. Periostracum not evident in specimens
examined. Nuclear whorls decollate in mature
specimens examined. Radula littorinid (2-1-1-1-2)
similar to L. unifasciata.
Animal darkly pigmented on surfaces of ten-
tacles snout and foot. Verge yellowish white in
color moderately short and thick with a basal flap
containing a penial gland and hyaline accessory
flagellum. Reproductive activity at its greatest
during warm months (Tel Aviv, Israel; Palant and
Fishelson, 1968), Probably oviparous and spawn-
ing pelagic capsules.
Measurements (mm.)—
length width no. whorls locality
18.2 11.1 5+ Ghana
14.0 9.0 5+ Ghana
12.8 7.9 4+ Ghana
11.3 7.4 4+ Valencia, Spain
10.8 6.9 5+ Port Elizabeth, So. Africa
10.5 ‘Tal 4+ Valencia, Spain
9.9 6.4 5+ Alexandria, Egypt
9.0 5.8 4+ Alexandria, Egypt
8.5 5.5 4+ Alexandria, Egypt
fos oye 3+ Alexandria, Egypt
Synonymy—
1791 Turbo punctatus Gmelin, Systema Naturae, ed 13, vol. 1,
p. 3597 (Senegalia); refers to Adanson, ‘Seneg’. Vol. 1,
p. 168, t. 12, fig 1, “Le Marnat”’; 1942, E. Fischer, et
al, Journal de Conchyliologie, vol. 85, p. 268, pl. 10,
figs 2a, 2b; Holotype in Museum d'Histoire Na-
turelle, Paris, 17 x 11 mm.
E
= -
y | | | |
| | [ | one |
4 | ee }—___|___ 4}_ 4 aaa
Plate 367. Geographical distribution of Littorina (Austrolit-
torina) punctata (Gmelin), in the Mediterranean, eastern At-
lantic and South Africa.
[05-359]
476 Austrolittorina
1826 Tricolia draparnauldii Audouin, 1826, Explication des
Planches, Savigny, Description de l Egypte, Histoire
Naturelle, vol. 1, pt. 4, Mollusques, p. 41, pl. 5, fig 19
(Egypt); Holotype in Museum d’Histoire Naturelle,
Paris.
1845 Litorina pulchella Dunker, Zeitschrift fiir Malako-
zoologie, vol. 2, p. 166 (Loanda [Luanda]); type in
Berlin Museum; 1847, Philippi, Abbildungen und
Beschreibungen Conchylien, vol. 2, p. 198, Litorina,
pl. 4 fig 8; 1853, Dunker, Index Moll. Guineam, p. 12,
pl. 2, figs 11-20.
1847 Litorina syriaca Philippi, Abbildungen und Beschrei-
bungen Conchylien. vol. 2, p. 165, Litorina pl. 3, figs
21-22 (Ora Syriae [Coast of Syria]); type in Berlin
Museum?
1858 Littorina guttata Reeve, Conchologia Iconica, vol. 10,
Littorina, pl. 14, fig 76 (St. Vincent, Cape de Verds);
not L. guttata Philippi [which is L. meleagris Potiez
and Michaud].
1932 Littorina perplexa Turton, The Marine Shells of Port
Alfred, South Africa p. 133, pl. 28, fig. 960. (Port
Alfred, South Africa; type in Oxford University Mu-
seum).
Types—Turbo punctatus Gmelin is based on
Adanson’s figure of “Le Marnat’’. The specimen
represented by that figure which is the holotype
of T. punctatus is in the Paris Museum (pl. 364
fig. 9, 10). The holotype of T. draparnauldii
Audouin is also in the Paris museum. The types
of L. pulchella Dunker and L. syriaca Philippi
may still be extant in the Berlin museum. Reeves
L. guttata is based on Philippi’s figure of a West
-
lauriius
AFRICANA
20° 60°
Plate 368. Geographical distribution of Littorina (Austrolit-
torina) africana (Philippi), in South Africa and Madagascar.
Joseph Rosewater
Littorinidae
Indian specimen. Reeve apparently believed that
it was actually the West African species, although
the name L. guttata Philippi is generally held to
be synonymous with the small spotted L. melea-
gris. Young individuals of L. punctata appear
superficially similar to L. meleagris and a com-
parison of the animal characters of the two is
needed. The holotype of Turton’s L. perplexa is
in the Oxford University Museum.
Records—SPAIN: Valencia (SMF; USNM). ITALY: Sicily
(USNM). COAST OF SYRIA: (SMF). LEBANON: Beirut
(USNM). ISRAEL: Tel Aviv (Palant and Fishelson, 1968).
EGYPT: Alexandria (USNM). ALGERIA: Oran (USNM).
MOROCCO: Melilla; Tangier; Casablanca (all USNM); Es-
saouira (SMF). SENEGAL: Goree (SMF; ZMC). CAPE
VERDE IDS: Mindelo, Sao Vicente (USNM). LIBERIA: Cape
Palmas. FERNANDO POO: Santa Isabel. CONGO REPUB-
LIC: Banana R. mouth (SMF). ANGOLA: Ambrizete (SMF);
20 km. S. of Luanda (USNM). SOUTH AFRICA: Langebaan,
Saldanha Bay; Dassen Id.; Table Bay; False Bay; Hermanus;
Breede R. mouth, St. Sebastian Bay; Still Bay; Mossell Bay;
Knysna (all K. H. Barnard, 1963); Port Elizabeth, Algoa Bay
(MCZ; USNM).
Littorina africana (Philippi, 1847)
(Pls. 364, 365)
Range—South Africa, from the west side of
Cape Peninsula, to southern Mozambique; Mada-
gascar.
Remarks—Littorina africana is distinct from,
but apparently closely related to, L. knysnaensis.
Both species occur on the south coast of Africa.
The appearance of its shell and its anatomy also
show similarities to other southern ocean species,
L. unifasciata, L. unifasciata antipodum and fer-
nandezensis and L. cincta of Australia and New
Zealand. The shell of L. africana differs from
these species in its development of comparatively
strong spiral sculpture (although some individu-
als appear almost smooth); anatomically its verge
is pigmented and the basal flap relatively more
pointed. Although Janus (1961) stated that the
South African species L. decollata Philippi is
distinct from L. africana the examination of large
series and a study of their ecology would be
necessary to settle questions regarding differ-
ences. Material examined during the present
study indicates that the two are identical. Tryon
(1887) considered this to be the case and acting as
first reviser chose L. africana as the senior syno-
nym in spite of the obvious page priority of the
name decollata.
Habitat—On rocks in the splash zone.
Description—Shell reaching 13.5 mm. (about .5
inch) in length, short-turbinate to subglobose in
shape, average obesity about .69 (10 specimens
range from .63-.77); older individuals only
[05-360]
November 30, 1970
moderately thick in structure, imperforate, usual-
ly developing a narrow, flattened crescent-shaped
area adjacent to columellar callous; sculptured
with closely-spaced, raised spiral threads of vary-
ing widths and irregular, sometimes coarse axial
lines of growth (sometimes without strong spiral
sculpture). External ground color grayish-white,
with a diffuse bluish gray band encircling the
body whorl and on anterior portions of spire
whorls. Aperture medium to dark brown with a
prominent white band near junction of outer lip
and columella and another often obscure band
high in aperture at junction of outer lip and body
whorl. Base somewhat flattened, separated from
upper part of body whorl by a low rather indis-
tinct keel at periphery. Whorls 3-5, rounded; spire
usually much eroded, considerably less than half
the length of shell, convex, produced at an angle
of about 66°. Aperture widely oval; outer lip
moderately thick, having its origin quite high on
body whorl, above keel, so that keel enters aper-
ture. Columella usually brown with a lighter
somewhat rimmed medial edge, moderately wide
and excavated anteriorly, and with a flattened to
excavated, brownish, crescent shaped area dis-
tally on the adjacent base. Suture impressed.
Sculpture consisting of ctosely-spaced, raised
spiral threads of varying widths; wider threads
usually separated by 2-4 narrow threads, disap-
pearing on base and on eroded spire, and some-
times appearing entirely smooth. Axial sculpture
consisting of occasionally coarse, irregular
growth lines. Operculum corneous, paucispiral.
Periostracum not evident in specimens examined.
Radula littorinid (2-1-1-1-2) central tooth some-
what narrow similar to L. unifasciata.
Animal darkly pigmented on surfaces of ten-
tacles, snout, foot and on edge of mantle. Verge
also darkly pigmented distally, fairly long and
club-shaped; having a basal flap which bears a
single penial gland containing a hyaline accesso-
ry flagellum. Nothing is known concerning the
reproduction and development of this species
although probably it is oviparous and spawns
pelagic capsule.
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 477
Measurements (mm.)—
length width no. whorls locality
13:5 8.4 4 Fort Dauphin,
Madagascar
10.7 7A 3+ South Africa
10.5 7.0 3+ Margate, Natal
9.9 6.7 4 Natal
9.9 tao 5 Natal
9.7 6.7 4 Natal
9.6 6.4 4 Isipingo, Natal
9.0 Bal 4 Umbhlali River, Natal
8.6 6.6 4 South Africa
8.2 5.9 3+ Inhaca Island,
Mozambique
re 5.0 3+ Margate, Natal
Synonymy—
1847 Litorina africana Philippi, Abbildungen und Beschrei-
bungen Conchylien, vol. 2, part 7, p. 199, Litorina, pl.
4, fig 10 (Caput Bonae Spei [Cape of Good Hope,
South Africa]; lectotype in Stuttgart Museum (N.H.),
catalogue number MT 106: 9.8 x 7.3 mm. (Janus,
1961); 1848, Krause, Die Sudafrikanischen Mollus-
ken, p. 102; 1858, Reeve, Conchologia Iconica, vol.
10, Littorina, pl. 8. figs 37 a, b.
1847 Litorina decollata Philippi, ibid., p. 196, pl. 4, fig 3 (Ora
Natal Africae [Coast of Natal, South Africa]); lectotype
in Stuttgart Museum (N.H.), catalogue number MT
107; 5.8 x 4.3 mm. (Janus, 1961); 1848, Krause, ibid.,
p. 102.
Types—Lectotypes of L. africana and L. decol-
lata are in the Staatlichen Museum fiir Natur-
kunde, Stuttgart, Germany, and were designated
from Krauss material by Janus (1961). Philippi
listed “Cape of Good Hope” as type locality for
L. africana and “coast of Natal” for decollata.
Krauss specified “Algoa Bay” for africana which
may be construed as a restriction of Philippi’s
type locality. Janus listed “Algoa Bay” and
“Natal” as the type localities of the lectotypes of
africana and decollata respectively.
Records—SOUTH AFRICA: W. Cape peninsula opposite
Simonstown; Robberg, Plettenberg Bay (both ANSP); Cape
Recife; Port Alfred (both MCZ); East London, mouth of
Nahoon River; Second Beach, Port St. Johns, Pondoland; Port
Edward, Natal; Margate, 11 mi. S. Port Shepstone, Natal (all
ANSP); Tiger Rocks, Isipingo, Natal (USNM; NMW); Dur-
ban, Natal (MCZ; USNM): mouth of Umhlali River, Natal.
MOZAMBIQUE: Inhaca Is. Delagoa Bay (both USNM).
MADAGASCAR: Pointe Ibanona, Fort Dauphin (MCZ).
[05-361 }
478 Austrolittorina
Joseph Rosewater
Littorinidae
Littorina knysnaensis (Philippi, 1847)
(Pls. 364, 365)
Range—South Africa, from Lambert’s Bay,
west coast to Natal.
Remarks—The “Knysna” littorina is quite dis-
tinctive and relatively easily distinguished from
other South African species. “Typical” specimens
exhibit a fairly prominent keel, flattened sculp-
ture and brownish and tan coloration with whit-
ish spots. The spots are most clearly observable
above and below the wide brown peripheral color
band. There is similarity, however, between the
verges of L. knysnaensis and L. africana. Where
differences in male anatomy are very slight, habi-
tat differences or physiological barriers may oper-
ate in preventing cross fertilization between the
species (see Palant and Fishelson, 1968). This
species occurs in part outside the range of other
South African Littorina, being the dominant
and/or only South African species on the west
coast and becoming rare farther north on the east
coast (Stephenson et al, 1940; Stephenson, 1947).
It is of course similar in part to L. punctata in its
spotted coloration, but differs in color pattern
distribution and in shell form.
The form L. africana tryphena Bartsch was
based on beach worn and anomalously high-
spired specimens whose appearance is very dif-
ferent. Nevertheless, tryphena is an absolute syn-
onym of knysnaensis.
Habitat—Intertidal on rocks and in crevices of
seawalls.
Description—Shell reaching 13.6 mm. (about
0.5 inch) in length, turbinate in shape, average
obesity about .64 (21 specimens range from .56-
.70); older individuals only moderately thick in
structure, imperforate, usually developing a nar-
row flattened, crescent-shaped area adjacent to
columellar callous; sculptured with spiral striae,
between which surface of shell is flattened, and
closely-spaced rather regular oblique axial lines
of growth; at times spiral and axial sculpture tend
to produce a reticulated pattern. External color
mahogany brown in a wide band above peripher-
al keel, lighter near suture and on the base, with
whitish spots and streaks especially on base.
Aperture dark-brown with a white band near
junction of outer lip and columella. Base distinct-
ly flattened, separated from upper part of body
whorl by a raised keel at periphery. Whorls 3-5,
somewhat flattened; spire usually eroded, less
than half the length of shell, convex, produced at
an angle of about 66°. Aperture oval; outer lip
moderately thick, having its origin quite high on
body whorl, above keel, so that keel enters aper-
ture. Columella brown, but sometimes cream-
colored, with a rather weakly rimmed medial
edge; moderately wide and excavated anteriorly,
and with a flattened to excavated, brownish to
cream colored crescent shaped area distally on the
adjacent base. Suture moderately impressed.
Sculpture consisting of about 9 spiral striae on
penultimate whorl of spire (surface of younger
whorls usually to worn to count) persisting onto
body whorl where 15-17 striae may be present
above keel, and 9-10 below. Surface of shell
between striae usually flat or only slightly raised.
Axial sculpture consisting of fine, regular oblique
axial lines of growth. Nuclear whorls about 3,
smooth, light brown; first post nuclear whorl
spirally sculptured. Operculum corneous, pauci-
spiral. Periostracum not evident in specimens ex-
amined. Radula littorinid (2-1-1-1-2), similar to L.
unifasciata; central tooth somewhat narrow.
Animal darkly pigmented on surfaces of ten-
tacles, snout, and foot. Verge may also be darkly
pigmented distally, moderately long and club
shaped; having a pointed basal flap with a penial
gland and a hyaline accessory flagellum; sperm
duct open, deeply folded. Nothing is known
concerning the reproduction and development of
this species although probably it is oviparous and
spawns pelagic capsules.
Measurements (mm.)—
length width no. whorls locality
13.6 7.7 4+ Algoa Bay, Cape Colony
13.0 8.0 4+ “South Africa”
12.5 8.0 4+ “South Africa”
12.2 6.9 5. Algoa Bay, Cape Colony
11.8 8.1 3+ “South Africa”
10.8 6.8 4 Muizenberg, Cape
Province
10.7 6.8 44+ Cape of Good Hope
9.7 6.2 4 Margate, Natal
9.6 6.6 3+ Knysna River,
Cape Province
8.7 6.0 4+ Algoa Bay, Cape Colony
8.6 5.8 5 Camps Bay,
Cape Colony
8.4 5.5 3 Port Elizabeth,
Cape Colony
8.0 5.3 5+ Capeland betw. Port
Alfred and Cape
Agulhas
7.6 4.6 4 Port Alfred,
Cape Colony
6.7 4.7 Q+ Cape of Good Hope
[05-362]
November 30, 1970
Synonymy—
1847 Litorina knysnaensis Philippi, Abbildungen und Besch-
reibungen Conchylien, vol. 2, p. 196, Litorina pl. 4,
fig 4, (Caput Bonae Spei ad regionem fluminis
Knysna [Cape of Good Hope, South Africa, near
Knysna River]); lectotype in Stuttgart Museum (NH),
catalogue number MT 108: 9.3 x 6.0 mm. (Janus,
1961); 1848, Krauss, Die Siidafrikanischen Mollus-
ken, p. 102.
1857 Littorina picea Reeve, Conchologie Iconica, vol. 10,
Littorina, pl. 15, fig. 83 (no locality given); lectotype
BM(NH) 1968320: 6.9 x 4.5 mm.
1915 Littorina africana tryphena Bartsch, United States Na-
tional Museum Bulletin 91, p. 120, pl. 38, fig. 6. (Port
Alfred, South Africa); holotype USNM 187091.
1932 Littorina rietensis W. H. Turton, the Marine Shells of
Port Alfred, S. Africa, p. 131, pl. 28, fig 948 (Port
Alfred, South Africa); type in Oxford University Muse-
um.
1932 Littorina kowiensis W. H. Turton, ibid., p. 132, pl. 28,
figs. [sic] 956 (Port Alfred, South Africa); type in
Oxford University Museum.
1932 Littorina africana pica ‘Reeve, W. H. Turton, ibid., p.
133 [invalid emendation of L. picea Reeve].
1932 Littorina indistincta W. H. Turton, ibid., p. 133, pl. 28,
fig 959 (Port Alfred, South Africa); type in Oxford
University Museum,
Types—A lectotype for Litorina knysnaensis
Philippi was designated by Janus (1961) from
among 8 syntypes in the Stuttgart Museum MT
108. A lectotype is here designated for Littorina
picea Reeve from among 3 syntypes in the British
Museum (NH): 1968320 (see pl. 364, figs. 19, 20).
The type locality for L. picea is here designated
as South Africa. The holotype (figured specimen)
of L. africana tryphena Bartsch is in the U.S.
National Museum: USNM 187091, and the para-
type originally associated with the holotype has
been recatalogued: USNM 664353. The types of
Littorina rietensis, L. kowiensis and L. indistincta
all described by Turton (1932) are in the Oxford
University Museum.
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Littorina 479
Records—SOUTH AFRICA: Lambert’s Bay (Stephenson, et
al, 1940); Saldanha Bay (ANSP); Dassen Id. (ZMA); Camps
Bay (USNM); Chapman’s Bay, W. of Cape Peninsula (ANSP);
Muizenberg (USNM); Gordons Bay, nr. Strand, False Bay
(ANSP; ZMA); Simonstown (ZMC); Simons Bay, E. Cape
Point (ANSP); Cape of Good Hope (USNM); N.E. of False
Bay; Buffels Bay, Cape Point; Onrust, 7 mi. W. Hermanus;
Hermanus; Cape Agulhas; Cape St. Blaize, Mossell Bay (all
ANSP): mouth of Knysna River, Cape Province (ex. Krauss,
MCZ); Beacon Isle, Plettenberg Bay (ANSP); Sea View, 16 mi.
W. of Port Elizabeth (MCZ); Beacon Point, Port Elizabeth
(ANSP); Algoa Bay, Port Elizabeth, Cape Colony (MCZ;
USNM); Capeland, Bushman’s River mouth (USNM): Kowie
R., Port Alfred (NMW); Port Alfred (ANSP; MCZ; USNM);
Esplanade, East London (USNM);: Coffee Bay, 1 mi. S. of
Umtata River; Port Edward, Natal (both ANSP); Second
Beach, Port St. John, Pondoland; Margate, 11 mi. S. of Port
Shepstone, Natal (both USNM).
20°
SEYCHELLES |S.
F “(§ Zanzibar
6 -
20 laurilius
Reumon
| is —-
40° o°
Plate 369. Geographical distribution of Littorina (Austrolit-
torina) knysnaensis (Philippi), in South Africa.
[05-363 ]
480
Austrolittorina
Joseph Rosewater
[ These occasional blank areas occur between
genera and subgenera to permit the insertion
of new material and future sections in their
proper systematic sequence. |
[05-364]
Littorinidae
November 30, 1970
Nodilittorina von Martens, 1897
Type: Littorina pyramidalis
Quoy and Gaimard, 1833
Abbott (1954) pointed out that Nodilittorina
von Martens should be considered a full genus,
and designated as its tvype-species L. pyramidalis
Quoy and Gaimard. He considered Nodilittorina
to include also the west Atlantic species L. tuber-
culata Menke with somewhat similar sculpture.
Habe (1956) pointed out a difference between
radulae of pyramidalis and tuberculata and on
that basis proposed a new subgenus for tuber-
culata, Echinolittorina, and at the same time sug-
gested that Nodilittorina s.s. is an Indo-Pacific
group. In the course of the present study, radulae
have been studied of the various Indo-Pacific
species thought to belong in Nodilittorina and all
have proven close in appearance to N. pyrami-
dalis.
In addition to the several Indo-Pacific species
belonging to Nodilittorina s.s., there are a num-
ber which because of sculptural differences are
here placed in the subgenus Granulilittorina
Habe and Kosuge, 1966. One species, N. natalen-
sis Philippi, because of its sculptural characters
resembles N. tuberculata. However, its radula is
not at all like Echinolittorina and, therefore, it is
maintained in the genus Nodilittorina s.s.
An anatomical character apparently common to
members ot Nodilittorina is the partial separation
of the penial gland from the basal enlargement of
the penis. This condition differs from Austrolit-
torina where the gland is fully incorporated in
the basal enlargement.
The most obvious character of Nodilittorina,
the “nodose” sculpture, is quite apparent in the
type-species, pyramidalis, in natalensis and
nodosa. It is not always so apparent in australis
with its highly variable sculpture. Nevertheless,
because of similarities in anatomy and the appar-
ent tendency to interbreed with nodosa it seems
obvious that australis must be placedin Nodilittor-
ina if current generic concepts are to be main-
tained.
Synonymy—
1897 Nodilittorina E. von Martens, in Weber’s Zoologische
Ergebnisse Einer Reise in Niederlandisch Ost-
Indien, vol. 4, pt. 1, p. 204; type-species by subse-
quent designation, Abbott, 1954, p. 451: Littorina
pyramidalis Quoy and Gaimard, 1833.
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 481
Subgenus Nodilittorina von Martens, 1897
Nodilittorina pyramidalis
subspecies pyramidalis (Quoy and Gaimard,
1833)
(Pls. 325, 326, 370, 371)
Range—West coast of India and southeast Asia
through the high islands of the Pacific to the
Marquesas.
Remarks—Nodilittorina pyramidalis is closely
related to several other species of the genus in the
Indo-Pacific, eastern Pacific and western Atlantic.
Present means of distinguishing these species are
based largely on differences in shell characters as
the various species do not differ detectably in
gross anatomy. When it is possible to examine
such features as comparative physiology, ecology
and the morphology of chromosomes additional
Plate 370. Figs. 1-9. Nodilittorina pyramidalis pyramidalis
Figs. 1,2. Holotype of Littorina pyramidalis Quoy and
Gaimard from Jervis Bay, Australia (MHNP, 23.7 x 13.7
mm.).
Figs. 3,4. Holotype of Littorina monilifera Eydoux and
Souleyet, from Touranne, Cochinchina (MHNP, 11.3 x 7
mm.).
Fig. 5. Lectotype of Littorina trochoides Gray (BM(NH)
87.4.26.1-8, about 10 x 7 mm.).
Figs. 6,7. Nodilittorina pyramidalis pyramidalis, from
Stradbroke Island, Queensland (USNM 684712, 18.5
12.8 mm.).
Figs. 8,9. The same from Koh Huyong, Similan Islands,
Thailand (USNM 661209, 8.6 x 5.2 mm.).
Figs. 10-13. Nodilittorina pyramidalis pascua Rosewater.
figs. 10,11. Holotype (USNM 679290, 13.1 x 8.8 mm.)
figs. 12,13. Paratype (USNM 679291, 12.6 x 7.8 mm.)
Both from Easter Island, Pacific Ocean.
[05-375]
482 Nodilittorina
Joseph Rosewater
Littorinidae
differences may be found. Arrangement of nodu-
lar sculpture has been found to be quite reliable
in the present study: N. pyramidalis usually
displays only two rows of white nodules on the
body whorl, one at the periphery and one above it
with only a single row on spire whorls (the
second being covered over by the succeeding
whorl); N. natalensis displays three rows on the
body whorl, the third row, usually slightly small-
er, being located at the suture, with 2-3 rows on
spire whorls and with a few raised spiral sculp-
tural cords running between the rows; N. sub-
nodosa shows a similar sculptural pattern, but it
is much reduced at times consisting of spiral rows
of weak granulations; the subspecies of N. py-
ramidalis (see next species) inhabiting Easter
Island is much like the nominate subspecies
except that the two rows of nodules on the body
whorl tend to coalesce producing axial bars rather
than nodules, and this may be noted on the spire
also; the west Atlantic species, N. tuberculata
Menke, is like N. natalensis in possessing 3 rows
of nodules on the body whorl and two on spire
whorls. This definite difference between N. py-
ramidalis and N. tuberculata contradicts the
statement by Abbott (1954) that the two are sib-
ling species as they are not strictly morphologi-
cally identical. The east Pacific species N. gala-
pagiensis Stearns has not been collected in suf-
ficient numbers to allow adequate understanding
of its variation. The latter species with three rows
of nodules on the body whorl appears more
closely related to the Atlantic and east African
species than to N. pyramidalis. Some young in-
dividuals of the east Atlantic species, miliaris
Quoy and Gaimard, have only two main rows of
nodules, but in later life develop multiple rows
which causes me to place them with the members
~
Plate 371. Nodilittorina pyramidalis pyramidalis Quoy and
Gaimard, from Koh Phi Phi, Thailand (USNM 661502).
Fig. A. Radula (upper scale is 0.05 mm.).
Fig. B. Penis; note partial separation of penial gland from
basal enlargement (lower scale is 2 mm.).
of |
: L f 2mm _y
of the subgenus Granulilittorina Habe and
Kosuge (see world species list).
Interestingly N. pyramidalis is one of those
species showing an affinity for high islands or
continental shores as indicated by its distribution
which skirts the Pacific atolls. Perhaps its prefer-
ence for high shore rocks limits the species to
coasts offering such a habitat.
On the coast of Queensland and New South
Wales, Australia, N. pyramidalis apparently
reaches its largest size (see measurements) and it
was from this region (Jervis Bay) that the species
was originally described. It is difficult to specify
the reason for this gigantism, but it may possibly
be related to a lack of optimal spawning tempera-
tures or to parasitic castration either of which
could prevent the onset of sexual maturity and
thus prolong the active growing period.
Habitat—Usually found on shore rocks con-
siderably above high tide line.
Description—Shell reaching 23.5 mm. (nearly |
inch) in length; but usually less than 12 mm.;
high cone or pyramidal in shape; average obesity
about .63 (43 specimens ranging from .55 to .70);
relatively thick in structure, imperforate, sculp-
tured with raised spiral cords and bearing rows of
raised nodules; microscopic sculpture, where not
worn away or otherwise obscured, consisting of
overall closely spaced wavy spiral threads. Axial
sculpture consisting of oblique growth lines. Ex-
ternal color, exclusive of nodules, dark reddish to
blackish brown, nodules usually white, but some-
times a lighter reddish brown. Aperture medium
to dark reddish brown, with a narrow yellowish
white band revolving inward from near anterior
junction of outer lip and columella; columella
also medium to dark reddish brown. Base some-
what flattened; periphery nodulated. Whorls 5-6,
rather flat-sided. Spire usually more than half the
length of shell, produced at an angle of from
45-60°. Aperture roundly oval to nearly diamond-
shaped; outer lip moderately thick; inner lip
(columella) flattened, moderately excavated, es-
pecially anteriorly where a broad siphonal trough
is directed at an angle of 45° to anterior-posterior
axis of shell. A flattened crescent shaped area on
base adjacent to columella callous. Suture distinct
and covering over lower row of nodules on spire
whorls. Predominant sculptural feature is double
row of white nodules on body whorl, each row
developing 11-15 nodules; nodules often crossed
by spiral cords; one row at periphery, the second
just above it; nodules usually arranged one above
the other, but sometimes out of phase; a single
[05-376]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 483
row of nodules on spire whorls; the peripheral
row on spire being covered over by next succeed-
ing whorl, but sometimes just visible above su-
ture. Nuclear whorls worn or decollate in all
specimens examined, remaining portions smooth;
postnuclear whorls nodulose. Operculum round-
ly oval, paucispiral. Radula littorinoid, formula
2-1-1-1-2; central tooth narrow, tricuspid; outer
marginal teeth with blunt enlargements laterally.
Animal darkly pigmented on upper surfaces of
tentacles, snout and foot. Verge relatively short
and thickened in preserved specimens, with an
enlargement near its base which is partly sepa-
rated from a penial gland bearing an accessory
flagellum; sperm groove deeply folded; distal end
of verge appears minutely papillose. Produces a
pelagic capsule having 6 spiral ridges, bearing a
single egg, and measuring 160 micra in diameter
(Tokioka, 1950; Habe, 1956).
Measurements (mm.)—
length width no. whorls locality
23.5 16.4 6 Lord Howe Id.
2h D: 14.2 6 Stradbroke Id.,
Queensland
21.8 14.5 6 Lord Howe Id.
19.3 Tay 6 Port Kembla, New
South Wales
18.4 12.3 5 Sydney, NSW,
1k7.3 12.1 6 Stradbroke Id.,
Queensland
13.4 8.0 5 “Marquesas”
11.6 7.0 5+ Berhala Id., Sandakan
No. Borneo
10.6 6.3 5 Pelaboean Ratoe, Java
7 5.9 5 Jamelo Bay, Luzon, P.1I.
9 6.1 5 Barrow Id., betw. Cape
Dupuy and Cape
Malouet, W. Australia
5.4 3.0 5 Guam, Marianas
Synonymy—
1791 Trochus nodulosus Gmelin, Systema Naturae, ed. 13,
vol. 1, part 6, p. 3582 (In Oceano australi, (et minor)
mari; Americam meridionalem); refers to Chemnitz,
Conchylien Cabinet, vol. 5, pl. 163, f. 1545, 1546; not
T. nodulosus Solander, 1766 in G. Brander, Fossilia
Hantoniensia, p. 10, pl. 6.
1817 Turbo trochiformis Dillwyn, A Descriptive Catalogue of
Recent Shells, vol. 2, p. 826 (Southern Ocean); refers
to Gmelin, T. nodulosus, p. 3582, and to Chemnitz,
ibid.; not T. trochiformis Brocchi, 1814.
1833 Littorina pyramidalis Quoy and Gaimard, Voyage de
L’Astrolabe, Zoologie, vol. 2, p. 482, pl. 33, figs.
12-15. (Jervis Bay [New South Wales] Australia);
holotype in Paris Museum, 23.7 x 13.7 mm.; 1954,
Abbott, Proceedings U.S. National Museum, vol. 103,
p. 456.
1839 Littorina trochoides Gray, the Zoology of Captain
Beechey’s Voyage—in His Majesty's Ship Blossom,
Mollusca, p. 140 (no locality); (lectotvype, BM(NH)
87.4.26.1-8, ca. 10 x 7 mm.)
1846 Litorina vilis ‘Menke’ Philippi, Abbildungen und Besch-
reibungen Conchylien, vol. 2, p. 145, Litorina, pl. 2,
fig 21 (no locality).
1847 Litorina malaccana Philippi, Abbildungen und Besch-
reibungen Conchylien, vol. 3, p. 15, Litorina, pl. 6,
fig 17 (Pulo Pinang).
1851 Litorina cecillei Philippi, Zeitschrift fur Malakozoo-
logie, 8th Jahr, p. 78 (Ryukyu Islands).
1852 Littorina monilifera Eydoux and Souleyet, Voyage sur la
Bonite, Zoologie, vol. 2, p. 559, pl. 31 figs 37-39
(Touranne, Cochinchine [Viet Nam]); holotype in
Paris Museum: 11.3 x 7 mm.
[1950 Littorina-capsula multistriata Tokioka, Publ. Seto
Marine Biological Laboratory vol. 1, no. 3, p. 151, fig
6, 2; non-binomial; is egg capsule of N. pyramidalis
fide Habe, 1956, Venus, vol. 19, no. 2, p. 121.]
Types—The lectotype of Trochus nodulosus
Gmelin and also of T. trochiformis Dillwyn is the
specimen figured by Chemnitz which may still be
extant in the Zoological Museum in Copenhagen,
Denmark. Holotypes of Littorina pyramidalis
Quoy and Gaimard and L. monilifera Eydoux and
Souleyet are in the Paris Museum. Types of
species described by Philippi: L. vilis, malaccana
and cecillei may be in the Berlin Museum.
Nomenclature—The excellent reviews of this
species by P. -H. Fischer (1967a, 1969) came to
my attention when the present study was well
advanced. Fischer’s decision that the oldest name
for this species is N. nodulosa (Gmelin, 1791) is
correct with the unavoidable qualification that
Trochus nodulosus Gmelin, 1791 is preoccupied
by Solander in G. Brander, 1766 (see Synonymy).
The next valid name, excluding T. trochiformis
Dillwyn, 1817, also preoccupied (Brocchi, 1814)
is Littorina pyramidalis Quoy and Gaimard,
1833. Although one may wish to clarify nomen-
clature by applying strictly the rules of priority,
one cannot dispense entirely with the effects of
homonymy. I agree otherwise almost entirely
with Fischer’s analysis of this species, with the
exception that I have not seen records from Mada-
gascar or the Red Sea and Persian Gulf areas and
can only suggest that Fischer's records from these
localities may be based on other species, such as
N. natalensis or subnodosa.
Records—INDIA: Bandra, N. of Bombay (USNM): Bombay
(MCZ); Vengurla, N. of Goa; Goa; Kumpta (Kumta), North
Kanara (all USNM); Cape Comorin; W. of Mandapam, Gulf of
Mannar; Rameswaram Island, Pamban, Palk Strait; Ramen
Point, W. side of Pamban Pass, btwn. Gulf of Mannar and
Palk Strait (all ANSP); Madras (MCZ, AMS, ANSP).
CEYLON: W. of Kankesanturai (ANSP); Galle (RNHL,
ANSP, USNM):; Columbo (MCZ, USNM); Merissa Village
(ANSP); Trincomalee (YPM). THAILAND: Ko Sindarar Nua
(Chance Island); Ko Huyong (South Island), Similan Islands;
Laem Phan-Pha, Ko Phuket, from around Phan-Pha point; Ko
Phi Phi (all USNM); Songkhla (MCZ); Ang Thong Id; Ko Tao;
Ko Maprao: Sriracha (all USNM): Ko Nom Sao, Chanthaburi
Province (MCZ); Ko Samet; Rayong; Lem Sing; Lem Ngob
(Ngop) (all USNM). MALAYSIA: Pulau Ular, Langkawi Is-
lands (USNM); Penang (MCZ); Malacca (MCZ); Pulau Anyut,
Malacca Strait, just S.E. of town of Malacca (USNM); Raffles
[05-377]
484 Nodilittorina
Joseph Rosewater
Littorinidae
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Plate 372. Geographical distribution of Nodilittorina pyrami-
dalis pyramidalis (Quoy and Gaimard) and of its subspecies,
N. pyramidalis pascua Rosewater.
Light, Singapore (ANSP, USNM); Sekudu Island, Strait of
Johore (USNM). VIETNAM: Ba Lang, North Annam
(USNM); Ile de la Table (AMS, MCZ). CHINA: Hong Kong
(ANSP, USNM); Amoy (Hsia-men); Spider Island, Foukien
Province (both USNM). JAPAN: Hachijo Island, 275 mi. S. of
Tokyo; Shirahama, Wakayama Pref. (both ANSP); Waki,
Satsuma (BPBM). RYUKYUS: Amami Oshima (ANSP); 1 mi.
N. of Shana Wan (USNM); Nago, both Okinawa (USNM,
ANSP, BPBM); Zenda, Kume (MCZ, USNM, ANSP, BPBM).
PHILIPPINE ISLANDS (Many localities; see map). NORTH
BORNEO: Sipitang; Sulok Island, Jesselton; Berhala Island,
Sandakan (all USNM); W. Marudu Bay (ANSP). INDONE-
SIA: Pulau Sebesi, Sunda Strait; Dijakarta (both RNHL):
Teluk Pelabuhan Ratu (MCZ, USNM); Welkomst Bay, Ban-
tam (USNM); Patjitan, all Java; Bali; Larantuka, Flores Island
(all RNHL). COCOS-KEELING IDS.: South Id. (Maes,
1967). AUSTRALIA: QUEENSLAND: Thursday Island,
(RNHL); Murray Island, both Torres Strait; No. VI Island
(both AMS): No. VIII Island, both Howick Islands (ANSP);
Lizard Island (AMS); Green Island (AMS, MCZ); Fitzroy
Island (AMS): Brook Island (ANSP); Palm Islands (AMS,
ANSP); Bay Rock, under clumps of dead coral and rock; Cape
Cleveland nr. Townsville (both USNM); Holbourne Island,
off Bowen; Hayman Island, Whitsundy Passage; Lindeman
Island (all AMS); Coppersmith Island, Smith Group (USNM);
Brampton Island; Coquet Island (both AMS); Yepoon; N.
Keppel Island, Yepoon (both AMS, MCZ); Keppel Bay; Heron
Island, Capricorn Group; Bustard Bay; Noosa Heads; Caloun-
dra (all AMS); Point Lookout, N.E. Stradbroke Island (ANSP,
AMS); Stradbroke Island (WAM). NEW SOUTH WALES:
Byron Bay; Woody Head (Wooded Bluft); Budgewoi Beach;
Toukley (all AMS); The Entrance (USNM); Putty Beach
(AMS); Wyargine Point, Middle Harbor, Sydney (AMS); Col-
laroy (USNM); Long Reef, N. of Manly (ANSP); Port Jackson
(ANSP, AMS, MCZ); Bottle and Glass Rocks, Sydney (ANSP);
Coogee Beach, Sydney (USNM); btwn. Gratlo Point and
Clontor, Sydney (AMS); La Perouse; Kurnell, both Botany
Bay (both MCZ); Bird Island, off Wollongong (AMS); Port
Kembla (USNM); Shellharbor (AMS); Jervis Bay (WAM,
MHNP): Sussex Inlet Reef (AMS); Twofold Bay (MCZ, AMS).
VICTORIA: Mallacoota (AMS). WESTERN AUSTRALIA:
Point Gregory, N.W. end of Peron Peninsula, Shark Bay;
below Quobba Light, N. of Carnarvon (both WAM); btwn,
Cape Duprey and Cape Malouet, Barrow Island (WAM,
USNM); mouth of False Cape Creek, La Grange Bay; 2 mi.
S.W. of jetty, Broome; James Price Point, 35 mi. N. of Broome
(all ANSP); Buccaneer Archipelago (AMS). NORTHERN
TERRITORY: Darwin (ANSP, USNM). NEW GUINEA: Ma-
nokwari; reef at S.E. entrance to Wooi Bay, Japen Island (both
ANSP); Yule Island (AMS, ANSP). LORD HOWE ISLAND:
(MCZ, AMS, NMW). NORFOLK ISLAND: (AMS). BONIN
ISLANDS: Port Lloyd; Ani Jima (both USNM):; Mukoshima
(ANSP, USNM). MARIANAS: Saipan (ANSP); Apra Bay,
Guam (USNM). SAMOA: Ofu Island; Muitre Point, Tau;
Fagamalo, Tau; Siulagi Point, Tau (all BPBM). TONGA:
Niuafo’ou (BPBM). MARQUESAS: Hona Nui, Ua Huka;
Hiva Oa; (both ANSP); Hanavave, Fatuhiva; Nukuhiva (both
USNM, ANSP); Eiao (ANSP); Ua Pou; Tahuata (both USNM).
Nodilittorina pyramidalis
new subspecies pascua Rosewater
(Pl. 370, figs. 10-13)
Range—Oeno, Pitcairn, Henderson and Easter
Islands, southeastern Pacific.
Remarks—As mentioned in connection with
the nominate subspecies, the concept of N. py-
ramidalis pascua as a subspecies inhabiting the
eastern-most extremities of the Indo-Pacific
faunal region appears quite valid. The subspecies
is distinguished primarily on the basis of its
nodular sculpture. The two rows of nodules pres-
ent on the whorls in pyramidalis s.s. have a
tendency to coalesce in pascua forming a series of
axial ridges.
The presence of Nodilittorina pyramidalis on
Easter Island was first mentioned by Dall (1908)
and it was later recorded by both Odhner (1922)
and Lamy (1936) based on separate collections.
None of these workers noted the differences be-
tween Easter Island populations and the species
elsewhere in the Indo-Pacific.
Habitat—On rocks above high tide line.
[05-378 ]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 485
Description—Very similar to N. pyramidalis
with exception that the double spiral row of white
nodules is replaced by a row of axial bars (result-
ing from coalition of juxtaposed nodules). Reach-
ing 15.3 mm. in length (about .6 inch); average
obesity about .64 (22 specimens range from .57 to
69). Usually only one row of small nodules on
base. Axial bars on spire whorls half covered by
undulating suture. Color more uniform than in
pyramidalis; generally overall bluish gray, axial
bars occasionally white or a dark brown; aperture
very dark brown, with a dark brown flattened
crescent shaped area on base adjacent to and
parallel with columella. Nuclear whorls about 2,
medium brown, smooth, shining; postnuclear
whorls with axial sculpture. Radula and anatomy
generally similar to N. pyramidalis pyramidalis.
Measurements (mm.)—
length width no. whorls locality
15.3 9.8 7+ All Easter Island
14.5 8.3 5+ Paratype
14.0 8.7 6+ Paratype
13.7 8.6 5+ Paratype
13.1 8.8 7 Holotype
12.8 8.2 7+ Paratype
12°3 7.8 6+ Paratype
10.1 6.2 6+ Paratype
9.3 6.3 6+ Paratype
8.9 5.8 6+ Paratype
Types—The holotype of Nodilittorina pyrami-
dalis pascua (a female; USNM 679290) and 24
paratypes (males and females; USNM 679291)
were collected in October, 1968, on Easter Island
by Pat McCoy. Additional paratypes from the
same source are in the Academy of Natural
Sciences, Philadelphia. The latin word pascua,
meaning “Easter” is used in the combination N.
pyramidalis pascua as a noun in apposition.
Records—Oeno Island (USNM); Pitcairn Island (BPBM;
USNM): Henderson Island (Smith, 1913); Easter Island
(USNM; ANSP).
Nodilittorina australis (Gray, 1826)
(Pls. 325, 373)
Range—Western Australia, from Esperance to
Vansittart Bay.
Remarks—Nodilittorina australis is endemic to
Western Australia. It is closely related to another
endemic species, Nodilittorina nodosa Gray, with
which it is found in at least part of its range living
in what appears to be the same or a very similar
ecologic niche on rocks low in the intertidal zone.
Comparison of gross anatomies and radulae show
no outstanding differences although the shells of
the two species are clearly distinct. Specimens
showing characters intermediate between N. aus-
tralis and nodosa found in the same population
with these species are judged to be hybrids (pl.
373, figs. 11-16). The observed similarities in the
anatomy of reproductive organs are believed to
permit successful inter-species copulation and
hybridization, although this phenomenon has not
been recognized often in Mollusca (see Boss,
1964). Hybrid specimens N. australis x N.
nodosa are fairly easily recognizable. They are on
the average more slender than either parent (.66
obesity versus .68) tend to show more typically
nodosa shell characters during early growth, but
in later growth take on more the appearance of
australis.
True australis may be recognized by its rather
globose shell, generally rough but not nodose
sculpture which is often wrinkled axially, light
tan to light violet colored aperture which has a
relatively broad white band revolving within it.
Spire sculpture is granulose but never nodulose.
In comparison, the shell of nodosa is pyramidal
with two rows of large whitish to orange nodules,
one row at the suture and one at the periphery of
the body whorl; on spire whorls there is one row
at the uppermost extremity and one at the lower-
most. Coloration of nodosa is dark brown, es-
pecially within the aperture, with a relatively
narrow revolving white band at its lower extremi-
ty. The shape of aperture in australis is roundly
oval, while in nodosa it is nearly diamond
shaped. Related to the shape of aperture is the
presence of a nodose keel in nodosa and the
virtual absence of a keel in australis. Hybrid
individuals exhibit a variety of characters gener-
ally intermediate between australis and nodosa.
Habitat—On rocks of the lower splash zone,
intertidal.
Description—Shell reaching 23.4 mm. (about
0.9 inch) in length, subglobose to subturbinate in
shape, average obesity about .68 (28 specimens
range from .63 to .75); relatively thick in struc-
ture, imperforate, usually developing a flattened,
crescent-shaped area adjacent to columellar cal-
lous; sculptured with rather heavy spiral cords
and often pronounced axial growth wrinkles. Ex-
ternal color grayish to yellowish white. Aperture
light yellowish tan to light violet, with a broad
white band near anterior junction of outer lip and
columella. Base hardly flattened with only a very
low keeling effect at periphery. Whorls 4-6, mod-
erately well rounded, especially the body whorl.
Spire less than half the length of shell, convex,
[05-379]
486 Nodilittorina
Joseph Rosewater
Littorinidae
produced at an angle of about 75°. Aperture oval;
outer lip moderately thick, inner lip concave,
thickened. Columella usually light violet to tan,
hardly excavated but with a flattened crescent-
shaped area on the adjacent base. Anterior junc-
tion of outer lip and columella projecting to form
a weak siphonal channel which occasionally
bears a series of very fine linear scratches. Suture
not deeply impressed. Spiral sculpture varying
from simple to moderately granulose, consisting
of from 5-8 raised spiral cords on spire whorls
and 18-20 on the body whorl; sculpture often
thrown into axial wrinkles by growth pattern,
either in part or over entire surface of shell—
giving a reticulated appearance. Entire surface
covered with closely spaced, fine, wavy, spiral
threads, most noticeable in grooves between spi-
ral cords; not detectable in worn specimens. Ir-
regular, fine axial growth lines also present. Oper-
culum corneous, only moderately thick, paucis-
piral. Periostracum not evident in specimens ex-
amined. Nuclear whorls about 3 in number, light
tan in color, smooth; first post-nuclear whorls
similarly colored but spirally sculptured, becom-
ing rapidly granulose. Radula littorinid (2-1-1-1-2)
central tooth very narrow (much like N. pyrami-
dalis).
Animal darkly pigmented on surfaces of ten-
tacles, snout and foot. Verge moderately short and
thick, yellowish white in color; having a bipartite
(mitten-shaped) basal flap the “thumb” ap-
pendage bearing a penial gland having a hyaline
accessory flagellum (similar to N. pyramidalis).
Nothing is known concerning reproduction and
development of this species, although it probably
is oviparous and spawns pelagic capsule.
Plate 373. Figs. 1-4, Nodilittorina australis (Gray, 1826).
Figs. 1,2. Neotype of Littorina australis Gray, from South
Mole, Fremantle, Western Australia (WAM 292-70, 15.1 x
11 mm.).
Figs. 3,4. An especially “rugose” specimen from Port Deni-
son, Western Australia (USNM 691677, 12.3 x 8.1 mm.).
Figs. 5-10. Nodilittorina nodosa (Gray, 1839).
Figs. 5,6. Lectotype, from north coast of Western Australia
(BM(NH) 87.4.26.10-12; 10.1 * 7.3 mm.).
Figs. 7,8. Specimen from Red Bluff, Kalbarri, near mouth of
Murchison River, Western Australia (USNM 691680, 13 x
8.9 mm.).
Figs. 9,10. A young specimen from the same locality (7.5 x
5.2 mm.). ;
Figs. 11-16. Supposed N. australia x nodosa hybrids; note
early “nodose” sculpture becoming more like australis in
figs. 11-14.
Figs. 11,12. from Vansittart Bay, northern Western Aus-
tralia (USNM 684714, 17.7 x 10.7 mm.).
Figs. 13,14. from Port Denison, Western Australia (USNM
691678, 14.4 x 9.2 mm.).
Figs. 15,16. Specimen from same locality as figs. 7-10,
possibly a young hybrid; note elongated nodules (8.2 x
5.7 mm.)
[05-380]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 487
Measurements (mm.)—
length width no. whorls locality
23.4 17.6 4+ “Australia”
19.0 12.4 6 All Red Bluff,
Kalbarri, West Australia
18.0 12.0 6
feo 11.0 o+
16.2 10.2 6
16.0 abi ies 4+
15.3 11.0 o+
15.1 10.2 5+
14.7 10.1 5+
13.8 9.5 5+
13.5 9.4 6
12.5 9.2 5+
T5 Tat 5+
10.7 hell 5
Synonymy—
1826 Littorina australis J. E. Gray in P. P. King, Narrative of a
Survey of the Intertropical and Western coasts of
Australia, vol. 2, Appendix B, p. 483 (type locality
here designated, South Mole [Arthur’s Head] mouth
of Swan River Fremantle, Western Australia.) Ne-
otype designated here: West Aust. Mus. 292-70; not
Littorina australis Gray, 1839, a Risella.
1843 Litorina rugosa Menke, Molluscorum Novae Hollandiae
Specimen pg. 9 (ad scopulos calcareous collis
Arthurshead, od ostium fluvii cygnorum [mouth of
Swan River, Fremantle, Australia]); 1844, Zeitschrift
fur Malakozoologie, Jahrgang 1844, p. 57 [synon-
ymizes L. rugosa with L. australis Gray, 1826].
Types—According to the notes given by Gray
(1826, ibid., p. 496) specimens of L. australis
were not among the species deposited by him in
the British Museum collections, nor were repre-
sentatives of this species found there during my
recent search for types of Littorinidae. It is fairly
certain, therefore, that “type-specimens of this
species and also that of its only other synonym, L.
rugosa Menke, may be considered lost. In order to
stabilize the concept of the species I consider it
necessary to designate a neotype (I.C.Z.N., Art.
10, 1961),
Neotype designation—Specimens of Littorina
australis Gray, 1826, were collected from rocks at
South Mole [Arthur’s Head], at the mouth of the
Swan River, Fremantle, Western Australia, by
B. R. Wilson and A. Paterson, February 9, 1968.
This is a locality from which specimens could
have been collected during Captain King’s survey
and it is here designated the type-locality for L.
australis. It is also the type-locality of L. rugosa
Menke. The specimen designated as neotype of L.
australis Gray, 1826, is catalogued as Western
Australian Museum 292-70 and is figured in pl.
373, figs. 1, 2. Additional specimens from the
neotype lot are catalogued as U.S.N.M. 679292.
Measurements of Neotype: length 15.1 mm.;
width 11.0 mm.
ee ee -
Rep orgie -
GIES
oop
YAva oe
-KEELING
DLLS Y
CS Se ee Se
oo° 120° 140°
Plate 374. Geographical distribution of Nodilittorina australis
(Gray), in Western Australia.
Records—AUSTRALIA: WESTERN AUSTRALIA: Esper-
ance (Hodgkin et al, 1966); Middleton Beach, nr. Albany
(WAM): Foul Bay (AMS); Margaret River (WAM); Augusta
(NMW); Yallingup, Cape Naturaliste (USNM, WAM); Ge-
ographe Bay; Cottesloe (both AMS); Cockburn Sound (WAM);
Rottnest Island (WAM, AMS): Garden Island, $.W. of Free-
mantle (WAM); Freemantle (USNM); Snag Island; Beagle
Islands (both WAM): Port Denison (USNM, WAM); Irwin
River (AMS); Abrolhos Islands (WAM); Pelsart Islands, Geel-
vink Chain, W. of Geraldton (AMS); Rat Island (WAM): Shark
Bay, Freycinet Estuary, Eagle Bluff; Denham, Shark Bay,
Peron Peninsula; S.E. Dirk Hartog Island (all WAM); Dirk
Hartog Island, Shark Bay (AMS); below Quobba Light, Blow
Holes, N. of Carnarvon (WAM); Red Bluff, Kalbarri, nr. mouth
of the Murchison River (USNM, WAM); Point Cloates nr.
Ningaloo (AMS); Mouth of Bigota Creek, Barrow Island;
btwn. Cape Dupuy and Cape Malouet, Barrow Island, (both
USNM, WAM); Vansittart Bay (USNM, AMS).
Nodilittorina nodosa (Gray, 1839)
(Pls. 325, 373)
Range—Western Australia, from the vicinity of
Geraldton northward [based on available rec-
ords].
Remarks—tThe failure of this species to appear
often in collections outside of Australian Muse-
ums is probably due in large part to its limited
distribution in Western Australia. It is also readi-
ly confused with N. australis because the two
species apparently hybridize and the hybrid
forms look like intermediates of a variational
continuum. The species is here considered to be
distinct although closely related to australis. It
may be recognized readily by its characteristic
diamond shaped outline and the two rows of
rather large whitish to reddish brown nodules on
the body whorl. See remarks under N. australis.
Habitat—On rocks of the splash zone, inter-
tidal, usually occurs with N. australis Gray.
[05-381]
488 Nodilittorina
Description—Shell reaching 13.3 mm. (about
0.5 inch) in length, with a diamond-shaped out-
line, average obesity about .68 (29 specimens
range from .64 to .72); moderately thick in struc-
ture, imperforate, usually developing a flattened
crescent-shaped area adjacent to columellar cal-
lous; with two rows of whitish to reddish brown
nodules on spire and body whorls. External color
very dark-brown between nodules. Aperture very
dark-brown with a relatively narrow white band
near anterior junction of outer lip and columella;
white spots on inner edge of outer lip mark inner
side of last nodules; columella usually lighter
brown than interior of aperture. Base somewhat
flattened, the effect accentuated by the large row
of nodules at periphery. Whorls 4-5, rather flat-
sided. Spire usually eroded, considerably less
than half the length of shell, convex, produced at
an angle of about 66°. Aperture diamond-shaped;
outer lip moderately thick; inner lip concave,
thickened. Columella usually light-brown to
light-violet, very shallowly excavated, with a flat-
tened crescent-shaped area distally on the ad-
jacent base. Anterior junction of outer lip and
columella project forming weak siphonal channel
which together with inner edge of outer lip occa-
sionally bear series of very fine linear scratches.
Suture not deeply impressed, but undulating
around nodules of preceding whorl. Spiral sculp-
ture of body whorl consisting of 2 rows, each
containing about 9 large nodules arranged at the
suture and periphery; penultimate whorl with
about 11 nodules in each of 2 rows at either
suture, the anterior row partly covered over by the
body whorl. Other spiral sculpture consisting of
rather low cords interspersed with fine wavy
white spiral threads. Axial sculpture consisting of
fine irregular growth lines and crevices resulting
190° 20° \40° 160°
Plate 375. Geographical distribution of Nodilittorina nodosa
(Gray), in Western Australia.
Joseph Rosewater
Littorinidae
from injury repair. Operculum corneous, rather
thin, paucispiral. Periostracum not evident in
specimens examined. Nuclear whorls eroded in
all specimens examined. Radula littorinid (2-1-1-
1-2) central tooth extremely narrow. Anatomy of
animal similar to that of N. australis.
Measurements (mm.)—
length width no. whorls locality
13.3 8.8 4+ All Red Bluff, Kalbarri,
Western Australia
12.5 8.4 3+
11.6 7.3 3+
10.9 12 4
10.0 6.5 3+
9.8 6.8 3+
8.8 5.9 4+
8.3 5.8 4+
6.4 4.4 3+
6.4 4.6 3+
Synonymy—
1839 Littorina nodosa Gray, the Zoology of Captain Beechey’s
Voyage—in His Majesty’s Ship Blossom, Mollusca,
p. 139 (no locality given; not figured [type locality
here designated: North coast of Western Australia]);
lectotype in British Museum (N. H.) 87.4.26.10-12;
10.1 x 7.3 mm. 1847, Philippi, Abbildungen und
Beschreibungen Conchylien, vol. 2, Litorina, p. 160,
pl. 3, fig. 7.
Types—The lectotype of Littorina nodosa
Gray, here designated, is in the British Museum
(NH) (see pl. 373, figs. 5, 6). It bears BM(NH)
catalogue number: 87.4.26.10-12; there are 2
paralectotypes under the same number. Measure-
ments of the lectotype are 10.1 x 7.3 mm.
Records—WESTERN AUSTRALIA: Port Denison (WAM;
USNM): North Island, Abrolhos Islands (WAM); Red Bluff,
Kalbarri (WAM; USNM); Quobba Point, 40 miles N. of
Carnarvon (ANSP; USNM); Northwest Cape (AMS); Vansit-
tart Bay (AMS; USNM).
[Nodilittorina australis x N. nodosa hybrids]
(Pl. 373, figs. 11-16)
Range—Coextensive with N. australis and N.
nodosa.
Remarks—See remarks under N. australis. In
shell characters hybrids appear intermediate be-
tween australis and nodosa. Their average shell
obesity is less than either parent: .66 compared
with about .68 (23 specimens range from .61-
.72). Gross anatomy of the two species is similar
and hybrids show no marked differences. Labora-
tory and field studies should be carried out to
determine whether introgressive hybridization is
taking place and other details of this phenome-
non.
[05-382]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 489
Habitat—On rocks of the splash zone, interti-
dal.
Measurements (mm.)—
length width no whorls locality
18.8 11.8 5 Port Dennison
18.2 11.6 4+ All Red Bluff,
Kalbarri, W. Australia
17.6 11.6 4+(eroded)
16.4 10.8 4+
15.8 10.5 4+
15.0 9.2 5+
14.4 9.8 4+
13-5 89 4+
13.0 9.2 34+
12.9 8.8 4+
12.0 8.7 3+
Synonymy—So far as can be determined no
scientific names have been established for these
hybrid forms. The combination N. rugosa x N.
nodosa has no validity or standing in zoological
nomenclature.
Nodilittorina natalensis (Philippi, 1847)
(Pl. 376, fig. 1-6)
Range—East Africa and Madagascar.
Remarks—Nodilittorina natalensis has a rather
narrow range in East Africa and the southwestern
Indian Ocean. It is replaced in south Asia and the
remainder of the Indo-Pacific by another some-
what similar appearing Nodilittorina, N. pyrami-
dalis. However, it may readily be distinguished
by the appearance of the nodular sculpture: nata-
lensis usually having at least three major spiral
rows of white nodules on the body whorl and 2 or
3 rows on the spire whorls, while pyramidalis has
two rows on the body whorl and one on the spire
whorls; natalensis also usually has a medium- to
light-brown spire apex, whereas the apex of pyra-
midalis is either eroded or not noticeably differ-
ent in color from the rest of the shell. In sculpture
natalensis is closer in appearance to the west
Atlantic N. (Echinolittorina) tuberculata than to
any Indo-Pacific species. The similarity does not
extend to the radula, however, that of natalensis
being much better developed than that of tuber-
culata whose radula appears to have suffered re-
duction in the numbers of cusps on the lateral and
marginal teeth and in the width of the central.
Habitat—Shore rocks above the splash zone.
Description—Shell reaching 14.5 mm. (about .6
inches) in length, pyramidal in shape; average
obesity about .64 (32 specimens range from .59 to
.69); relatively thick in structure, imperforate,
20° 40° 60° 80°
20°
ns
7
MALDIVE » >
is. 7g
SEYCHELL:S IS.
CHAGOsS |”
is. 24
D Mauracs
i
Reumon
NATALENSIS
60° 80°
Plate 377. Geographical distribution of Nodilittorina natalen-
sis (Philippi), in the western Indian Ocean.
Plate 376. Nodilittorina (Nodilittorina) natalensis (Philippi,
1847). Note 3 rows of nodules at and above periphery.
Figs. 1,2. Lectotype, from Coast of Natal, South Africa
(Stuttgart Museum (NH): MT 109, 12.2 x 8.5 mm.; photo
from Janus, 1961, Stuttgarter Beitrage zur Naturkunde,
no. 70, pl. 3, figs. 7,8).
Figs. 3,4. Specimen from Point Ibanona, Fort Dauphin,
southeast Madagascar (USNM 679281, 13.1 x 7.7 mm.).
Figs. 5,6. Possible paralectotypes “ex Krause’, from Natal
(Senckenberg Museum, 11.5 x 7.9 mm.).
[05-383]
490 Nodilittorina
Joseph Rosewater
sculptured with raised spiral cords and bearing
rows of raised nodules; microscopic sculpture,
where not worn away or obscured, consisting of
overall, closely-spaced spiral threads. Axial
sculpture consisting of fine lines of growth. Ex-
ternal color, exclusive of nodules and apex, dark
blackish brown; nodules usually grayish to yel-
lowish white. Aperture medium to dark reddish
brown, with a narrow white band revolving in-
ward from near anterior junction of outer lip and
columella, in some specimens there is a wider
light colored band near posterior junction of outer
lip and columella. Apex light to medium brown.
Base flattened with nodulose cords. Whorls 5-7,
very slightly rounded. Spire usually more than
half the length of shell, occasionally nearly coe-
qual with length of aperture, produced at an angle
of from 53-64°. Aperture roundly oval—
subquadrate; outer lip moderately thick; inner lip
(columella) moderately flattened and excavated,
especially anteriorly where a broad siphonal
trough is directed at an angle of 45° to anterior-
posterior axis of shell; a flattened, crescent-
shaped area on base adjacent to columella callous.
Suture rather indistinct. Predominant sculptural
feature: three rows of white nodules on body
whorl, each row developing 13-15 nodules often
crossed by spiral cords; one row at periphery, the
second just above it, the third just below suture;
nodules usually arranged one above the other, but
sometimes out of phase; occasionally one or two
accessory nodulated rows may develop from spi-
ral cords between primary rows, especially high
on whorl. Two to three nodule rows appear on
spire whorls. Spiral cords on base often moder-
ately nodulose. In an exceedingly well-preserved
specimen from Chango Island, Zanzibar (ANSP)
with nuclear whorls intact, the latter consist of
about two smooth, light brown volutions; first
postnuclear whorl also smooth, but succeeding
whorls spirally striate and becoming nodulose.
Operculum thin, chitinous, light-brown, oval,
paucispiral. Radula littorinid, formula 2-1-1-1-2;
central tooth very narrow, tricuspid, the outer
cusps held high above center cusp and close to
body of tooth.
Littorinidae
Animal darkly pigmented on anterior upper
surface of head, and also tentacles, snout and
foot. Verge relatively short and thick in preserved
specimens, with an enlargement near its base
which is partly separated from a penial gland
bearing an accessory flagellum; sperm groove
deeply folded. Reproduction unknown; probably
oviparous, producing pelagic capsule.
Measurements (mm.)—
length width no. whorls locality
14.3 9.1 6+ Fort Dauphin,
Madagascar
13.7 8.4 6+ Fort Dauphin,
Madagascar
13.2 7.8 7+ Fort Dauphin,
Madagascar
12.2 7.3 7+ Fort Dauphin,
Madagascar
11.7 7.2 6+ Coast of Natal (ex
Krauss
10.5 al 6+ Coast of Natal (ex
Krauss
9.6 5.9 6+ Faty, Madagascar
8.9 6.1 6+ Coast of Natal (ex
Krauss
8.3 4.9 7+ Faty, Madagascar
7.4 4.9 6+ Coast of Natal (ex
Krauss
6.8 4.5 4 Inhaca Island,
Mozambique
Synonymy—
1847 Litorina natalensis Philippi, Abbildungen und Beschrei-
bungen Conchylien vol. 2, p. 160, Litorina pl. 3, fig 4.
(Coast of Natal, South Africa); lectotype in Stittgart
Museum, MT 109: 12.2 x 8.5 mm. (Janus, 1961);
1848 Krauss, Die Sudafrikanischen Mollusken, p.
102.
Types—Janus (1961) designated a lectotype for
L. natalensis Philippi from specimens collected
by Krauss and deposited in the Staatlichen Muse-
um fiir Naturkunde in Stuttgart (pl. 376 figs. 1, 2).
Records—SOUTH AFRICA: Cape of Good Hope (ANSP);
Second Beach, Port Saint Johns, Pondoland; Margate, 11 mi.
S. of Port Shepstone, Natal; Port Edward, Natal; Tiger Rocks,
Isipingo, Natal (all ANSP); Durban, Natal; mouth of the
Umhlali River, Natal (both MCZ). MOZAMBIQUE: Inhaca
Island, Delagoa Bay, (USNM, ANSP). TANZANIA: Chango
(Prison) Island, W. Zanzibar (ANSP). ADEN _ PRO-
TECTORATE: Conquest Bay (USNM). MADAGASCAR:
Faty (Ifatz), 13 mi. N. of Tulear; Anako, 20!/2 mi. S. of Tulear;
Pointe Ibanona, Port Dauphin; Flacourt, Fort Dauphin;
S.W. shore Le aux Nates, S. of Ile Ste. Marie; 2.5 mi. N.E. of
Pointe D’Antsiraikiraiky, N.W. Ile Ste. Marie (all MCZ).
[05-384 ]
November 30, 1970
Subgenus Granulilittorina
Habe and Kosuge, 1966
Type: Granulilittorina millegrana (Philippi, 1848)
The subgenus Granulilittorina forms a con-
venient group for those rather globose Nodilit-
torina having multiple rows of low granular
sculpture as contrasted with the members of
Nodilittorina s.s. which are more pyramidal in
shape and usually exhibit only one to three rows
of larger nodules. The radulae and other anatomi-
cal details appear grossly similar in the two
subgenera.
Synonymy—
1966 Granulilittorina Habe and Kosuge, Shells of the World in
Colour, vol. 2, The Tropical Pacific, p. 20, pl. 6, fig.
13; Venus, vol. 24, no. 4, pp. 313, 328; type-species by
monotypy Granulilittorina philippiana Habe and
Kosuge, 1966 [= Nodilittorina (Granulilittorina) mil-
legrana (Philippi, 1848)].
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 491
Nodilittorina millegrana (Philippi, 1848)
(Pls. 326, 378, 379, 380)
Range—From the Red Sea and western Indian
Ocean islands to the western Pacific where it
occurs sporadically as far east as the Marshall
Islands.
Remarks—Better known by several of its syno-
nyms L. millegrana is an extremely variable
species throughout its range in the Indo-Pacific.
Much of the variation may be ecologically in-
fluenced. A form exhibiting reticulate sculpture,
possibly representing the doubtful species L.
reticulata Anton, occurs on Indian Ocean Islands
and in the Pacific has been found sporadically in
the Philippines, and as far eastward as Eniwetok,
Marshall islands, apparently showing a prefer-
ence for islands. Axially and spirally striped color
forms occur more or less randomly throughout
the species range, but perhaps appear more com-
monly in the Philippines, East Indies and Aus-
tralia. Variation in shell granulation is consider-
Plate 378. Nodilittorina (Granulilittorina) millegrana (Phi-
lippi, 1848), showing shell variations.
Figs. 1,2. Noumea, New Caledonia (USNM 679730, 11.8 x
7.6 mm.).
Figs. 3,4. Near Jesselton, North Borneo (USNM 658369, 8.0
x 5.3 mm.).
Figs. 5,10. Barrow Id., Western Australia (USNM 691690,
9.2 x 6.4 mm.).
Figs. 6,7. Tsutsu, Tsushima Id., Nagasaki, Japan (USNM
679181, 8.7 x 5.5 mm.).
Figs. 8,9. Point de Galle, southwest Ceylon (USNM
672392, 12.2 x 8.5 mm.).
Figs. 11,12. Goa, India (USNM 442974, 10 x 6.7 mm.).
Figs. 13,14. Eniwetok Id., Eniwetok Atoll, Marshall Ids.
(USNM 679731, 9.3 x 6.2 mm.).
[05-395]
492 Granulilittorina
able. Its limits are at present considered to in-
clude forms ranging from smooth to rather
heavily granulose. Constant characters include
the generally globose appearance, the fact that the
aperture is usually more than half the length of
shell and the more or less dependable presence of
some degree of granulation. This is the Littorina
granularis “Gray of authors” (non Gray). As
shown by Yen (1942), the species granularis has
as its type an Atlantic specimen and is probably
synonymous with WN. miliaris (Quoy and
Gaimard) from the Eastern Atlantic, rather than
Tectarius muricatus (Linné) as suggested by Yen
(see pl. 325).
Habitat—On shore rocks above high tide line.
Description—Shell reaching 13.7 mm. (about
0.5 inch) in length, subglobose to subturbinate in
shape; average obesity about .67 (56 specimens
range from .59 to .76); moderately thick in struc-
ture, imperforate; surface occasionally smooth,
but more often bearing raised, granulose spiral
cords on spire and body whorls; overall micro-
scopic sculpture where not worn away or other-
wise obscured consisting of many fine wavy spi-
ral threads most easily observable between
primary granulose spiral cords. Axial sculpture
usually limited to irregular axial lines of growth,
occasionally, especially in island populations,
granulations are in phase axially and reticulate
appearance results. External color variable, rang-
ing from overall grayish white to spotted and
axially and spirally striped with yellow to reddish
brown markings. Aperture medium to dark
brown, often the former lighter color with stripes
or spots of the darker color, usually with a white
band revolving inward from near anterior junc-
tion of outer lip and columella; columella also
medium to dark brown. Base not greatly flat-
tened; periphery not markedly differing in its
contour from rest of body whorl. Whorls 5-7,
well-rounded. Spire usually less than half the
length of shell, produced at an angle of from
about 68-73°. Aperture widely oval; outer lip
moderately thick, occasionally slightly flaring,
inner lip (columella) strongly developed, broad,
thick, occasionally bulging posteriorly, shallowly
excavated; with a flattened crescent shaped area
on base adjacent to columella callous, at times
approaching a columella chink. Suture well im-
pressed. Predominant sculptural feature, 9-10
spiral cords bearing granulations on body whorl
and 4-5 cords on spire whorls; granulation-
bearing cords usually interspersed with non bear-
ing cords, the latter often weaker; granulations
Joseph Rosewater
Littorinidae
often subdued and some shells virtually smooth.
Nuclear whorls brown to black, smooth and shin-
ing; postnuclear whorls becoming rapidly granu-
lose and usually having a single dark spiral color
band. Operculum paucispiral, brown, oval. Radu-
la littorinid, formula 2-1-1-1-2; central tooth
only moderately narrow, tricuspid.
Animal darkly pigmented on upper surface of
tentacle, snout and foot. Verge long and slender,
with an enlargement near its base partly separated
from a penial gland containing an accessory fla-
gellum; sperm groove deeply folded. Produces a
pelagic egg capsule, having 3 tiers and an undu-
late border (see Tokioka and Habe, 1953.)
Measurements (mm.)—
length width no. whorls locality
13.7 9.2 5 Goa, India
13.7 8.1 5 Bombay, India
12.3 8.6 4+ Pt. de Galle, Ceylon
11.3 7.3 5 Pondicherry, India
10.7 7.2 4 Goa, India
9.7 6.3 5 Koh Phi Phi, Thailand
8.4 5.4 6 Souillac, Mauritius
8.1 5.6 5 Hong Kong
7.8 5.4 4 Barrow Island,
Western Australia
6.7 4.2 4 Réunion
5.9 4.5 5 Keppel Bay,
Queensland
5.5 3.6 4 Koh Huyong, Thailand
Synonymy—
—Littorina granularis ‘Gray’ of Authors, non Gray, 1839
[Gray’s holotype, BM(NH) 87.4.26.9 is an eastern
Atlantic species, Nodilittorina miliaris (Quoy and
Gaimard].
1847 Litorina picta marmorata Philippi, Abbildungen und
Beschreibungen Conchylien, vol. 2, Litorina, p. 167,
pl. 3, fig. 26 [lectotype figure] (provincia Ilocos
borealis insulae Luzon); not L. marmorata Pfeiffer,
1839.
1847 (Sept.) Litorina ventricosa Philippi, ibid., vol. 3, Litorina
p. 51, pl. 6, fig 19 [lectotype figure] (Pulo Pinang
[Penang Island, Malaysia]); not L. scabra ventricosa
Philippi, 1847 (April).
1848 Litorina millegrana Philippi, ibid., p. 65, pl. 7, fig. 15
[lectotype figure] (Red Sea), original measurements
about 13 x 11 mm.
1852 Littorina radiata Eydoux and Souleyet, Voyage sur la
Bonite, vol. 2, p. 562, pl. 31 figs. 46, 47 (Touranne,
Cochinchine [Viet Nam]); lectotype BM(NH) 54.
7.24.389, 11.5 x 7.6 mm.
1857 Littorina novaezelandiae Reeve, Conchologia Iconica,
vol. 10, Littorina pl. 14, fig. 74 (New Zealand [Trin-
comali, Ceylon, here corrected]). lectotype BM(NH)
1966124, 13.2 x 9.0 mm. designated by Biggs,
1966, Journal of Conchology, vol. 26, no. 2, p. 138,
pl. 7, fig 5.
1857 Littorina granocostata Reeve, ibid., pl. 15, fig 79 (Bris-
bane Water [New South Wales] Australia); lectotype
BM(NH) 1968318, 6.3 x 4.4 mm.
21859 Littorina vidua Gould, Proceedings of the Boston
Society of Natural History, vol. 7, p. 138. (Ousima [=
Amami-o-shima, Ryukyu Islands, according to John-
son, 1964]); type lost.
[05-396]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 493
i
Plate 379. Nodilittorina (Granulilittorina) millegrana (Phi-
lippi, 1848).
Fig. A. Radula of specimen from Aden (USNM 679341;
lower scale is 0.05 mm.).
Fig. B. Penis of specimen from Sekudu Id, near Singapore,
Malaysia (USNM 660756; upper scale is 1 mm.).
1876 Littorina melanacme E. A. Smith, Journal of the Linnean
Society of London, Zoology, vol. 12, p. 552, pl. 30,
fig. 21 (San Christoval, Solomon Islands); Holotype
BM(NH) 76.1.10.67, 9.6 x 6.1 mm.
1885 Littorina erronea Nevill, Hand List of Mollusca in the
Indian Museum, part 2, p. 152 (Balapiti, Ceylon);
new name for Littorina novaezealandiae Reeve, 1857.
1887 Littorina granicostata E. A. Smith, Proceedings of the
Zoological Society of London for 1887, part 3, p. 519,
fig. 2, (Christmas Island, Indian Ocean); Holotype
BM(NH) 1968358, 10.4 x 7.2 mm.; is Littorina insu-
laris E. A. Smith, 1889.
1889 Littorina insularis E. A. Smith, Proceedings of the
Zoological Society of London for 1888, part 4, p. 536;
new name for L. granicostata E. A. Smith, 1887.
1897 Littorina ventricosa strubelli von Martens, Zoologische
Ergebnisse Einer Reise in Niederlandisch Ost-
Indien, vol. 4, part 1, p. 208 (Krakatau, Indonesia);
type in Zoological Museum Amsterdam ?; refers to L.
pusilla, and to Philippi, “Abbild. Neuer Conch. vol.
2, p. 164, pl. 4, fig 15 [pl. 3, fig 23?] and to Kuster,
Conchylien-Cabinet, vol. 2, pt. 9, p. 11, p. 1 fig.
20-22; 1925 Jutting, Treubia Batavia, vol. 6, p. 142
[not a new variety of Jutting’s as indicated by H. B.
Preston, Zoological Record, vol. 62, Mollusca, p. 54].
1915 Littorina eudeli Sowerby; Annals and Magazine of
Natural History, series 8, vol. 16, No. 93, p. 167, pl.
10, fig 5 (Pondicherry, India); helotype BM(NH)
1919, 12.31.33,12x 74mm. Lch;
1936 Littorina chaoi T-c Yen, Notes de Malacologie chinoise,
vol. 1, fas. 3, Musee Heude, Shanghai, p. 3 (Pok-hoy
{Gulf of Tonkin] China); 1937, ibid., fas. 4, figs 2, 2a,
2b; holotype in Museum Heude?
Plate 380. Type specimens of synonyms of Nodilittorina
(Granulilittorina) millegrana Philippi, 1848).
Fig. 1. Lectotype figure of Litorina millegrana Philippi,
from the Red Sea (from Abbildungen and Beschreibungen
Conchylien, vol. 3, Litorina, pl. 7, fig. 15, about 13 x 11
min.).
Figs. 2,3. Lectotype of Littorina radiata Eydoux and Soul-
eyet, from Touranne, Viet Nam (BM(NH) 54.7.24.389;
Lib X76 mm:):
Figs. 4,5. Lectotype of Littorina novaezelandiae Reeve, a
smooth specimen from Trincomali, Ceylon (BM(NH)
1966124; 13.2 x 9 mm.).
Figs. 6,7. Holotype of Littorina melanacme E. A. Smith,
from San Cristobal, Solomon Ids. (BM(NH) 76.1.10.67;
9.6 x 6.1 mm.).
Figs. 8,9. Holotype of Littorina insularis E. A. Smith [is
also holotype of L. granicostata Smith] from Christmas
Id; Indian Ocean (BM(NH) 1968358, 10.4 x 7.2 mm.).
Fig. 10. Holotype of Littorina urieli Biggs, from Eilat,
Israel (BM(NH) 1966121, 7.3 x 4.5 mm.).
Figs. 11,12. Lectotype of Littorina granocostata Reeve,
from Brisbane Water, Australia (BM(NH) 1968318, 6.3 x
4.4 mm.). lee Fes
Figs. 13,14. Holotype of Littorina eudeli Sowerby, from
Pondicherry, India (BM(NH) 1919. 12.31.33, 12 x 7.4
mm.).
1953 [Littorina-capsula hagruma Tokioka and Habe, Publica-
tions of the Seto Marine Biological Laboratory, vol. 3,
no. 1, pp. 55, 56 (Tanabe Bay, Japan); Habe, 1956,
Venus, vol. 19, no. 2, pp. 117-121, fig. B; non-
binomial].
1966 Granulilittorina philippiana Habe and Kosuge, Venus,
vol. 24, no. 4, pp. 313, 328, [figured in] Habe and
Kosuge, 1966, Shells of the World in Colour,
Hoikusha, vol. 2, p. 20, pl. 6, fig 13 (Goza, Shima
Peninsula, Honshu, Japan); Holotype in National
Science Museum, Tokyo, 6.8 x 5.2 mm.
1966 Littorina urieli Biggs, Journal of Conchology, vol. 26, p.
137, pl. 7, figs 1, 2 (Bilat, Gulf of Eilat, northern end
Gulf of Aqaba, Israel) Holotype, BM(NH) 1966121,
7.4 X 5.0 mm.
Types—The types of species described by Phi-
lippi in Abbildungen und Beschreibungen Con-
chylien should be in the Berlin Museum, al-
though I have not been able to confirm their
presence. Until this is possible Philippi’s figures
[05-397]
494 Granulilittorina
Joseph Rosewater
Littorinidae
must be considered accurate representations of
the species, and are here held to be the lectotypes.
It is interesting to note that the lectotype
(figured specimen) of L. radiata Eydoux and
Souleyet is in the British Museum (N.H.), while
only two paralectotypes were found at the Paris
Museum. Although it would be expected that the
‘Bonite’ collections would be in the Paris Muse-
um exclusively, a large number of the mollusk
types of Eydoux and Souleyet were bequeathed
by Souleyet to the British Museum (NH) (see
Gray, 1855).
A lectotype for Littorina novaezelandiae Reeve
was designated by Biggs (1966) and the type
locality is here corrected from New Zealand to
Trincomali, Ceylon. Other types of Reeve, Smith,
Sowerby and Biggs are in the British Museum
(NH).
The type of Littorina chaoi Yen may be in the
Museum Heude, Shanghai. The holotype of Gran-
ulilittorina philippiana Habe and Kosuge is in
the National Science Museum, Tokyo.
Nomenclature—The problem of selecting a
proper name for this species from among possible
synonyms is compounded because of its variabil-
ity, the fact that somewhat similar species occur
elsewhere in the oceans, and because some early
workers neglected to cite type localities with their
species descriptions. The earliest valid name ap-
Plate 381. Figs. 1,2. Holotype of Littorina miliaris Quoy and
Gaimard, 1833, from Ascension Id., Atlantic Ocean (MHNP,
14.6 x 9.4 mm.).
Figs. 3,4. Holotype of Littorina granularis Gray, 1839 [east-
ern Atlantic] (BM(NH) 87.4.26.9, 14.2 x 10 mm.).
pears to be L. millegrana Philippi, 1848. (see
synonymy). The tamiliar name L. granularis
Gray, 1839, has as its holotype a worn specimen
which was interpreted by Yen (1942) as being
Tectarius muricatus Linné. I believe it to be
actually a worn specimen of Nodilittorina miliar-
is (Quoy and Gaimard) a west African species.
There is also a possibility that L. reticulata
Anton, 1839, may figure in the competition for the
oldest name for this species. However, reticulata
like granularis Gray, has no type locality and the
only illustration, that given by Philippi and said
to be of Anton’s type shows a specimen which is
impossible to identify with certainty. Littorina
picta marmorata, usually considered to be a vari-
ant of the Hawaiian species picta Philippi, but
actually described from the Philippines, and L.
ventricosa Philippi, are both junior homonyms
and therefore fail to qualify as valid names,
leaving millegrana as the next available taxon.
Records—RED SEA: Eilat, Israel (G. Frankel Coll.). GULF
OF ADEN: Aden (ZMC). MADAGASCAR: S.W. Shore Ile des
Nattes, S. of Ile Ste. Marie; Ste. Luce, S.E. Madagascar (both
USNM). INDIAN OCEAN ISLANDS: E.. of Souillac
(Savanne R.), Mauritius; Caves Point, W. Mauritius; Réunion;
Dunidu Id., N. Male Id., Maldives; Fadiffolu Atoll, (all
USNM). INDIA: Bandra, N. of Bombay; Bombay; Goa; Cape
Comorin (all USNM); Pondicherry (USNM, AMS). CEYLON:
S. Shore Fort Frederick, Trincomalee; Pt. de Galle (both
USNM). THAILAND: Laem Phan-Pha, Koh Phuket; Koh
Huyong, Similan Ids.; Koh Phi Phi (all USNM):; Songkla
(MCZ); Koh Maprao; Koh Tao (both USNM); Prachaup Khiri
Khan (MCZ); Koh Nom Sao, Chantaburi Province (MCZ);
Koh Sichang; Koh Chala; Rayong; Koh Kut (all USNM).
MALAYSIA: Penang (MCZ); Batu Ferringgi, Penang (MCZ);
Pulau Ular, Langkawi; Pulau Jerak, W. of Sembilan Ids.;
Pulau Anyut, Malacca Str.; Pulau Besar, Malacca Str. (all
USNM); Raffles Light, Singapore (USNM, ANSP); Sekudu
Id., Johore Str. (USNM); CHINA: Hong Kong; Big Wave Bay,
Hong Kong. VIETNAM: (USNM). RYUKYUS: Odomari, Oki-
nawa, (USNM). JAPAN: Tsutsu, Tsushima Id., Nagasaki
Pref., Kyushu (USNM); Miura Peninsula and Goza, Shima
Peninsula, Honshu (both Habe and Kosuge, 1966). PHILIP-
PINES: Santo Domingo, Batan; Jamelo Bay; S. Shore, Fort
Mills, Cavite; Maricaban Id. (all USNM); Corregidor Id.
(USNM; ANSP): Mariveles; Grand Is., Subic Bay; Bolinao
Bay; Lingayen, all Luzon (all USNM); Marinduque; Pt. Naso,
Panay; Zamboanga, Mindanao; Balabac; Bucas Id.; Jolo;
Tumindao (all USNM). INDONESIA: Keledjitan, Bantam,
Java; Pelabuhan Ratu, Preager, Java; Krakatoa (ZMC); Sulok
Id., Jesselton, N. Borneo; Morotai, Halmahera Group, East
Indies, Moluccas (all USNM). COCOS-KEELING ATOLLS:
(ANSP). AUSTRALIA: QUEENSLAND: Lizard Id. (AMS);
Los Isles (USNM); Halfmoon Bay, nr. Cairns; Fitzroy Id.;
Palm Id. (all AMS); Bay Rock (USNM); Royal Seaforth, nr.
Lindeman Id.; Brampton Id.; Heron Id., Capricorn Grp. (all
AMS); Keppel Bay (USNM); North Keppel Id.; Barney Pt.,
Port Curtis; Lady Elliot Id.; Bundaberg; Point Vernon, Her-
vey Bay; Caloundra (all AMS); Cape Cleveland (USNM).
NEW SOUTH WALES: nr. Wollongong. WESTERN AUS-
TRALIA: btwn. Cape Dupuy and Cape Malouet, Barrow Id.;
mouth Bigota Creek, W. side Barrow Id.; Airport Beach,
Barrow Id.; Broome (all USNM). MELANESIA: S.E. En-
trance Wooi Bay, Japen Id., West Irian (ANSP); Karkar Id.,
North-East New Guinea; Matupi Isl. Rabaul, Bismarks;
Makira Hbr., San Cristobal, Solomons; Taden Reef, N. of
Heinghene, New Caledonia (all AMS); N. of Touho, New
Caledonia; E. of Nani Id. (both USNM); Noumea, New
Caledonia (AMS, USNM); Suva; Irvines, nr. Malaqereqere,
both VitiLevu, Fiji (both USNM). PALAU IDS: Melekeiok,
Babelthaup Id. (USNM). MARSHALL IDS.: Eniwetok
(USNM).
[05-398 ]
November 30, 1970
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Plate 382. Geographical distribution of Nodilittorina (Granu-
lilittorina) millegrana (Philippi), in the Indian Ocean and
western Pacific, and of N. (G.) picta and N. (G.) cinerea
(Pease), in Hawaii and the Marquesas Islands respectively.
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 495
Nodilittorina subnodosa (Philippi, 1847)
(Pl. 383, figs. 1-10)
Range—Red Sea and Persian Gulf areas.
Remarks—Nodilittorina subnodosa is an ex-
ceedingly variable species within its rather nar-
row area of habitation in the Red Sea and Persian
Gulf. The variation is expressed particularly in
the degree of shell granulation, different in-
dividuals ranging from distinctly nodulose to
hardly sculptured. One character apparently rare
in Nodilittorina, but present in this species, is the
white columella, which serves to distinguish
subnodosa from natalensis, its nearest geographic
relative. The relationships of subnodosa are
somewhat obscure. Certain individuals from Per-
sian Gulf populations resemble in part the east-
ern Atlantic N. miliaris Quoy and Gaimard (Rev.
H. E. Biggs, personal communication, 1968). I
consider that the resemblance is superficial, and
that the two are quite distinct. I believe, however,
that subnodosa may possibly share a common
ancestry with N. miliaris. The two Persian Gulf
populations examined indicate that N. sub-
nodosa is more weakly sculptured and may reach
a larger size there than in the Red Sea. However,
considerable additional comparative material is
needed to prove this trend in shell morphology.
Habitat—Shore rocks above high tide line.
Description—Shell reaching 13.8 mm. (about
My inch) in length, pyramidal to pyramidal-
turbinate in shape; average obesity about .66 (38
specimens range from .60 - .70); relatively thick in
structure, usually imperforate with some larger
and older individuals developing small umbilical
opening; sculpture varying from weakly granu-
lose to only moderately nodulose; microscopic
sculpture where not worn away consisting of fine
closely spaced, wavy spiral threads; axial sculp-
ture consisting of fine, irregular lines of growth.
External color, exclusive of nodules and apex,
yellowish to pinkish white, occasionally spotted
dark brown between nodules; nodules white.
Aperture reddish orange to dark brown, with a
narrow white band revolving inward from near
anterior junction of outer lip and columella;
white band obscure in some specimens. Dark
apertural coloration usually ending at anterior
junction of outer lip and columella; columella
white. Apex light to medium brown. Base moder-
ately to considerably flattened, sculptured with
spiral, granulose cords. Whorls 5-7, hardly to
moderately rounded. Spire usually more than half
the length of shell, produced at an angle of from
[05-399]
496 Granulilittorina
55-62°. Aperture oval, outer lip moderately thick,
tending to be slightly shouldered at the suture;
inner lip (columella) thick and moderately flat-
tened; shallowly excavated anteriorly; a flattened,
crescent shaped area on base adjacent to columel-
la and an umbilical opening occasionally present
in older individuals. Suture indistinct only in
more nodulose specimens. Predominant sculp-
ture: three spiral rows of medium sized nodules
above periphery on body whorl, often with addi-
tional rows of smaller nodules between principal
rows; usually 1-3 principal rows of nodules on
spire whorls with rows of smaller granulations
between. Sculpture in some specimens severely
muted, consisting only of spiral rows of granula-
tions showing little or no size separation. Nuclear
whorls worn or decollate in all specimens ex-
amined, light brown in color; postnuclear whorls
spirally nodulose. Operculum thin, chitinous,
light brown, paucispiral.
Anatomy grossly similar to N. natalensis. Re-
production unknown.
Records—RED SEA: (MCZ); N.E. Museri Id., Dahlak
Archipelago, Ethiopia (RNHL); Eilat, Gulf of Aqaba
(USNM). PERSIAN GULF: Bushire, Iran (ZMC); near Abu
Dhabi, Trucial Coast (USNM).
Joseph Rosewater
Littorinidae
Types—tThe location of the type specimens of
Litorina subnodosa Philippi is not definitely
known, although it is probable that they are in the
Berlin Museum. Of the two specimens repre-
sented in Philippi’s illustration, the larger, his fig.
9, is here designated as the lectotype (see pl. 383,
fe. 2)
Measurements (mm.)—
length width no. whorls locality
13.8 8.8 5+ Abu Dhabi,
Trucial Coast
12.8 8.3 5+ Abu Dhabi,
Trucial Coast
12.3 8.6 5+ Abu Dhabi,
Trucial Coast
11.9 8.0 6 Abu Dhabi,
Trucial Coast
1.1 7.2 5+ Abu Dhabi,
Trucial Coast
10.4 7.0 4+ Abu Dhabi,
Trucial Coast
9.7 6.8 4+ Abu Dhabi,
Trucial Coast
9.0 6.3 4+ Abu Dhabi,
Trucial Coast
8.4 5.3 7 Eilat, Gulf of Aqaba
tee 4.3 6 Eilat, Gulf of Aqaba
6.2 4.2 6 Eilat, Gulf of Aqaba
5.2 3.5 5+ Eilat, Gulf of Aqaba
Synonymy—
1847 Litorina subnodosa Philippi, Abbildungen und Besch-
reibungen Conchylien, vol. 2, p. 161, Litorina, pl. 3,
figs 8, 9 (Red Sea).
Plate 383. Nodilittorina (Granulilittorina) subnodosa. (Phi-
lippi, 1847)
Fig. 1. Lectotype figure of Litorina subnodosa Philippi,
from the Red Sea (from Abbildungen and Beschreibungen
Conchylien, vol. 2, Litorina, pl. 3, fig. 9, about 19 x 12
mm.).
Figs. 2,3. Specimens from an old collection from Gulf ot
Suez (USNM 23233, 14.5 x 9.6 mm.).
Fig. 4. Rather smooth specimen from near Abu Dhabi,
Trucial Oman Coast, Persian Gulf (USNM 669124, 11.8 x
7.8 mm.).
Figs. 5,6. Bushire, Iran, eastern Persian Gulf (USNM
679285, 8.2 x 5.3 mm.).
Figs. 7-10. Strongly nodulose to only moderately granulose
specimens from Eilat, Gulf of Aqaba, Red Sea (USNM
671239, 7,9: 8.3 x 5.2 mm.; 8,10: 7.5 x 4.9 mm.).
[05-400]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 497
20° 40° 60° 80°
eos ae ae tec — —
20°
MALDIVE » ©
1s. -
|
SEYCHELLES IS.
‘ 1 |
Plate 383a. Geographical distribution of Nodilittorina (Granu-
lilittorina) subnodosa (Philippi) in the Red Sea and Persian
Gulf.
Nodilittorina leucosticta
subspecies leucosticta (Philippi, 1847)
(Pl. 384, figs. 1-5)
Range—India and Ceylon.
Remarks—It is extremely difficult to quantify
the differences between G. leucosticta and G.
millegrana and on occasion it has seemed impos-
sible to maintain them as separate species. Never-
theless populations of leucosticta occur together
with millegrana in which the individuals be-
longing to each species are clearly separable.
Outside the supposed range of leucosticta, no
Plate 384. Nodilittorina (Granulilittorina) leucosticta and sub-
species.
Figs. 1-5. N. leucosticta leucosticta (Philippi).
Fig. 1. Lectotype figure of Litorina leucosticta Philippi
[Bombay, India] (from Abbildungen and Beschreibungen
Conchylien, vol. 2, Litorina pl. 3, fig. 11, about 8 x 6
mm.).
Figs. 2,3. Pointe de Galle, southwestern Ceylon (USNM
672389, 9.5 x 6.2 mm.).
Figs. 4,5. Goa, India (USNM 442970, 13.1 x 7.8 mm.).
Figs. 6-13. N. leucosticta biangulata (von Martens).
~ Figs. 6,7. Holotype of Littorina biangulata von Martens,
from Benkulen [southwest] Sumatra (ZMA, 11 x 7.1
mm.).
Figs. 8,9. Pulau Nias, Mentawai Islands. southwest of
Sumatra (USNM 654444, 6.6 x 4.1 mm.).
Figs. 10,11. Mouth Hienghene River, Hienghene Area,
New Caledonia (USNM 637368, 10 x 6.3 mm.).
Figs. 12,13. West tip Corregidor Id, Luzon, Philippines
(USNM 637360, 9 x 5.5 mm.).
Figs. 14-19. N. leucosticta feejeensis (Reeve).
Figs. 14,15. Lectotype of Littorina feejeensis Reeve,
(BM(NH) 1968319, 7.1 x 5 mm.).
Figs. 16,17. Ovatoa, northwestern Vanua Levu, Fiji (USNM
694772, 7.1 x 4.7 mm.).
Figs. 18,19. Tutuila, Samoa (USNM 488720, 7.2 x 4.7 mm.).
[05-401]
498 Granulilittorina
such mixing of phenotypes occurs. It is, therefore,
fairly apparent that G. leucosticta is a valid
species with more restricted range than mill-
egrana and that the two are clearly separable on
shell morphology alone.
The shell of leucosticta averages more slender
than millegrana (obesity .65 vs. .67). The single
most. striking feature in Jleucosticta is the
shouldered appearance of the body whorl in
mature individuals. The pattern of hardly raised
white nodules on the spiral cords is also distinc-
tive as these are usually interrupted with brown
spots; in some populations the shell ground color
is dark gray to brown, and here the white semi-
nodulated markings are very prominent. The
markings are always less pronounced, however,
than in N. pyramidalis. The aperture in leucostic-
ta usually is proportionately more than half the
length of shell except in large and apparently
anomalous individuals which are not rare in
collections.
The subspecies leucosticta biangulata von
Martens and lI. feejeensis Reeve replace leucostic-
ta s.s. in the East Indies and western Melanesia
and in the Pacific Islands respectively. Differ-
ences between the subspecies are of a consider-
ably qualitative nature and involve a shift in
emphasis of intensity of expressed sculpture and
coloration. If these taxa are to be considered
subspecies rather than distinct species or even
portions of a variational continuum, one must
refer to specimens from the several geographical
areas cited and to the descriptions of the various
morphological peculiarities of each. In such cases
species and subspecies concepts are considerably
strained.
Habitat—Intertidal on rocks.
Description—Shell reaching 16.2 mm. (about
°/s inch) in length, turbinate to elongate oval in
shape; average obesity about .65 (22 specimens
range from .59 to .70); only moderately thick in
structure; imperforate; surface sculptured with
rather low and closely spaced spiral cords 2-4 of
which, usually three, on body whorl, are stronger
than the rest; cords bearing low white, often
elongate nodules separated by brown interspaces;
shell shouldered about #/3 length of body whorl
from suture to first strong cord; details of color
and sculpture becoming obscured in larger in-
dividuals; overall microscopic sculpture where
not worn away or otherwise obscured consisting
of fine wavy spiral threads. Axial sculpture lim-
ited to irregular, often closely spaced lines of
growth. External color variable, ranging from
Joseph Rosewater
Littorinidae
grayish white base color to a dark gray or brown,
usually with the low white nodules offering dis-
tinct contrast. Aperture medium to dark brown,
with a white band revolving inward near anterior
junction of outer lip and columella, with occa-
sional fainter bands posteriorly; columella usual-
ly lighter colored than rest of aperture. Base
slightly flattened; periphery often marked by po-
sition of one of stronger spiral cords. Whorls 5-6;
spire whorls only moderately rounded; body
whorl rather flatsided at its center and shoul-
dered. Spire usually less than half the length
of shell, produced at an angle of from about
62-70°. Aperture elongate oval, outer lip only
moderately thick; inner lip strongly produced,
rather straight, only occasionally slightly bulging
posteriorly, with a flattened crescent shaped area
on base adjacent to columella callous, Suture well
impressed. Body whorl subtly shouldered; pre-
dominant sculptural feature: about three well
spaced, low spiral cords on body whorl; cords
with low white, sometimes elongate nodules,
separated by brown color spots; sculpture char-
acteristics often worn away or not evident in older
individuals. When present, nuclear whorls about
1*/2-2, light brown, smooth; postnuclear whorls
becoming darker brown and spirally sculptured.
Operculum paucispiral, brown, oval. Radula lit-
torinoid, central tooth only moderately narrow
tricuspid.
Animal darkly pigmented on upper surfaces of
tentacles, snout and foot. Verge of the Nodilit-
torina type, with a basal enlargement and _par-
tially separated penial gland with an accessory
flagellum; main portion of verge relatively long
and slender, with an open deeply folded sperm
duct along its posterior-medial edge. Nothing is
known concerning the reproduction and develop-
ment of this species, although it is suspected to
involve a pelagic capsule as yet undescribed.
Measurements (mm.)—
length width no. whorls locality
16.2 9.8 5+ All Bombay, India
15.1 9.3 5+
14.7 8.7 4+
13S 8.3 4+
11.0 7.1 3+
10.7 7.2 4+
9.8 6.8 4+
8.7 6.1 3+
7.9 5.2 4+
6.6 4.5 3+
Synonymy—
1847 Litorina leucosticta Philippi, Abbildungen und Besch-
reibungen Conchylien, vol. 2, Litorina, p. 162, pl. 3,
[05-402]
November 30, 1970 INDO-PACIFIC MOLLUSCA, vol. 2, no. 11 Nodilittorina 499
80° 100° 120° 140° 160° 180° 160° lac
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TOKELAU IS 1
Plate 385. Geographical distribution of Nodilittorina (Granu-
lilittorina) leucosticta leucosticta (Philippi) and of its other
two subspecies, N. (G.) leucosticta biangulata (von Martens)
and N. (G.) leucosticta feejeensis (Reeve).
fig 11 [lectotype figure; original measurement about 8
x 6 mm.] (no locality given; Bombay, India, here
designated); lectotype may be in Berlin Museum.
1866 Melaraphe subgranosa Dunker, Verhandlungen der
Kaiserlich-Koniglichen zoologisch-botanischen Ge-
sellschaft in Wien, Jahrgang 1866, vol. 16, p. 913
(Madras [India]); type may be in Berlin Museum;
1867, Frauenfeld, Reise der Osterreichischen Fre-
gatte Novara, Zoologischen Theil, vol. 2, part 3,
Mollusken, p. 9, pl. 1, figs. 10 a, b [not fig 10 ¢ which
G. millegrana Philippi].
1887 Litorina leucostica ‘Philippi’ Tryon, Manual of Con-
chology, vol. 9, p. 299 [error for L. leucosticta Phi-
lippi, 1847].
Types—Philippi's figure of leucosticta (the lec-
totype figure) depicts a young specimen, a phe-
notype apparantly quite common in India and
Ceylon, although this particular oval white spot-
ted form has not been noted in collections from
elsewhere. Proportions of the subspecies G. leu-
costicta biangulata are quite different. The lecto-
type figures of Frauenfeld of subgranosa Dunker,
show the species in its mature form.
Records—INDIA: Bandra, N. of Bombay; Bombay; Goa;
Cape Comorin; Tuticorin; Mandapam Camp. CEYLON:
Point de Galle; Columbo (all USNM),
Nodilittorina leucosticta subspecies
biangulata (von Martens, 1897)
(Pl. 384, figs. 6-13)
Range—East Indies and [?] western Melanesia.
Remarks—Littorina leucosticta biangulata von
Martens differs from the nominate subspecies in
having typically only two strong spiral sculptural
cords on the body whorl. Other sculpture is much
suppressed. Shells of this subspecies also tend to
be more high spired and therefore the tendency
for the spire to be less than half the length of the
shell is somewhat reduced. Shells reach 12.4 mm.
(about .5 inch) in length; average obesity about
.62 (29 specimens range from .56-.65). In the
representatives of populations available to me for
study, the conservatism in spiral sculpture exhib-
ited by the type is shown in only a few speci-
mens—most develop 3 moderately strong spiral
cords bearing subdued white nodules, more like
leucosticta s.s., but with the above-mentioned
narrower profile (see illustrations, also remarks
under leucosticta leucosticta).
Habitat—On intertidal rocks.
Measurements (mm.)—
length width no. whorls locality
12.4 7.0 6 Touho, New Caledonia
11.0 7.1 6 Holotype: Benkulen
Sumatra
10.4 6.8 5+ Jesselton, No. Borneo.
10.1 6.0 54+ Touho, New Caledonia
9.7 6.0 4+ Touho, New Caledonia
8.6 ee, 5+ Touho, New Caledonia
7.8 D0 6 Jesselton, No. Borneo
7.1 4.4 6 Corregidor Id., Luzon,
Philippines
6.7 4.2 5+ Pulau Nias, Sumatra
5.7 3.6 6 Pulau Nias, Sumatra
Synonymy—
1897 Littorina biangulata von Martens, in Max Weber, Zoo-
2 =
logische Ergebnisse einer Reise in Niederlandisch
[05-403]
500 Granulilittorina
Ost-Indien, vol. 4, part 1, p. 209, pl. 9, fig 26
(Benkulen [Bengkulu, SW Sumatra]); holotype in
Zoologisch Museum Amsterdam, 11 x 7.1 mm.
Records—PHILIPPINES: West tip Corregidor Id. Luzon
(ANSP, USNM); Santo Domingo, Batan Id., Batanes Group
(USNM). EAST INDIES: Jesselton, N. Borneo; Pulau Nias,
(both USNM); Bengkulu, both S.W. Sumatra (Von Martens,
1897; ZMA): Kahatola Id., S. Loloda Ids. Halmahera, Moluc-
cas; Pelabuhan Ratu, Preager, Java (both USNM); NEW
CALEDONIA: mouth Hienghene R., Hienghene area; 18 km.
N. of Touho, N.W. New Caledonia (both USNM).
Nodilittorina leucosticta subspecies
feejeensis (Reeve, 1857)
(PI. 384, figs. 14-19)
Range—Pacific Islands, exclusive of Western
Melanesia and Hawaii.
Remarks—According to available records Gran-
ulilittorina leucosticta feejeensis replaces leuco-
sticta s.s. and l. biangulata in the Pacific Islands.
It usually has a smaller, more compact shell and
spiral sculpture and axial color striping are often
strongly expressed, although from the present
appearance of Reeve’s type, this would not be
expected. The type of L. feejeensis Reeve has
apparently undergone some corrosion of sculp-
ture in over 100 years in the BM(NH) collection.
Shells reach 12.7 mm. (about .5 inch) in length;
average obesity about .64 (23 specimens range
from .61-.72). Most strikingly colorful popula-
tions appear to occur in Samoa and Tonga where
zigzag dark axial markings stand out against the
lighter ground color of the shell (see pl. 384, figs.
18, 19). The general shape of the shell and charac-
ter of sculpture and coloration clearly relate this
species most closely to leucosticta and I. bian-
gulata.
Habitat—Shore rocks.
Measurements (mm.)—
length width no. whorls locality
ay 7.8 5+ Ofu, Manu’a, Samoa
11.0 7.5 5 Ofu, Manu’a, Samoa
10.3 6.4 5 Tongatapu
9.4 6.0 5 Lifu, Loyalties
8.7 5.4 6 Arue, Tahiti
8.2 5.7 5 Niuafou
7.6 5.5 4 Makatea, Tuamotus
7.6 5.1 5 Kermadecs
6.5 4.5 5 Niuafou
5.6 4.0 4 Tutuila, Samoa
Synonymy—
1857 Littorina feejeensis Reeve, Conchologia Iconica, vol. 10,
Littorina, pl. 15, figs 82 a, b. (Feejee [sic] ids.);
lectotvpe BM(NH) 1968319: 7.1 x 5 mm.
1871 Litorina vitiensis ‘Reeve’ von Martens, Donum Bis-
marckianum, p. 40; emendation of L. feejeensis
Reeve, 1857.
1871 Litorina (Melaraphe) vitensis Dunker, Malakozoo-
logische Blatter, vol. 18, p. 150 (Hab. ad insulas
Vitenses).
Joseph Rosewater
Littorinidae
1885 Littorina plena var. vitiensis ‘Dunker’ Nevill, Hand-List
of Mollusca in the Indian Museum, Calcutta, Part 2,
p. 139 (Tahiti); emendation of L. vitensis Dunker,
1871.
1885 Littorina miliaris var. fijiensis ‘Reeve’ Nevill, ibid., p.
154 (Tahiti); emendation of L. feejeensis Reeve, 1857.
1951 Nodilittorina miliaris ‘Quoy and Gaimard’ Habe, Illus-
trated Catalogue of Japanese Shells, vol. 1, no. 14, p.
92, pl. 14, fig 5; not N. miliaris (Quoy and Gaimard)
which is Eastern Atlantic species and not from As-
cension Id., Pacific = Ponape, Caroline Ids.
Records—(All USNM except where noted). BONIN IS-
LANDS: Port Lloyd, Chichi Jima; Ani Jima. MELANESIA:
Vanikoro, Santa Cruz Group (AMS); Lifu, Loyalties. FIJI: W.
side Koro Levu Id. Taveuni. KERMADECS: Raoul, (AMS).
SAMOA: Pago Pago, Tutuila; Tafuna, Tutuila; Ofu, Manu’a
Group. TONGA: Niuafou; Niutoua, fringing reef, Tongatapu;
Lualea reef, nr. Fatuma, Tongatapu. SOCIETIES: Puputeai,
nr. Mt. Taharaa, Arue, Tahiti; Mt. Taharaa, Dist. of Mahina,
Tahiti; N. coast of Tahiti. TUAMOTUS: Makatea (USNM,
ANSP); N. of Tamao Hbr., Makatea; Maiai Id., Tikahau.
Nodilittorina exigua (Dunker, 1860)
(PI. 386, figs. 1-6)
Range—Coasts of China and Southern Japan.
Remarks—Although G. exigua is apparently
closely related to G. millegrana Philippi, an ex-
amination of representatives of Japanese and
China coast populations reveals a species that is
generally more strongly sculptured, lacks the
color striping, has a stronger siphonal trough and
more deeply excavated columella than mill-
egrana. Habe (1951) synonymized exigua under
Nodilittorina granularis (Gray) but it has been
demonstrated that granularis is a synonym of the
Atlantic species, N. miliaris (Quoy and Gaimard).
(see remarks under millegrana).
Habitat—Shore rocks.
Description—Shell reaching 12.5 mm. (about
0.5 inch) in length, high-spired globose to turbi-
nate; average obesity about .68 (22 specimens
range from .64 to .72); moderately thick in struc-
ture, imperforate; spiral sculpture usually prom-
inent and consisting of closely spaced raised
spiral cords which are alternately granulose and
smooth; granulose cords considerably more con-
spicuous; axial sculpture limited to occasionally
coarse oblique lines of growth. External color
fairly constant, usually a rather dirty yellowish to
grayish white. Aperture medium to dark reddish
brown; inner edge of outer lip often lighter
colored with spots of brown marking sculptural
furrows; with a broad white band revolving in-
ward from near anterior junction of outer lip and
columella; columella usually lighter than aper-
ture, occasionally as dark. Base only moderately
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 501
Plate 386. Figs. 1-6. Nodilittorina (Granulilittorina) exigua
(Dunker)
Figs. 1,2. Lectotype figures of Litorina exigua Dunker,
from Japan (from Dunker, 1861, Mollusca Japonica, pl. 2,
fig. 3).
Figs. 3,4. Matsu Shima, Sea of Japan (USNM 601593, 9.8 »
6.8 mm.).
Figs. 5,6. Imaizumi, Kagoshima Bay, Japan (USNM
363708, 8.9 x 6 mm.).
Figs. 7-12. Nodilittorina (Graulilittorina) picta (Philippi).
Figs. 7,8. Lectotype of Littorina picta Philippi (BM(NH)
1968324, 9.8 x 6 mm.).
flattened; periphery not carinate. Whorls 4-5,
rounded. Spire less than half the length of shell,
produced at an angle of from 68-78°. Aperture
oval; outer lip moderately thick, usually wrinkled
at edge; inner lip (columella) moderately well
developed, rather deeply excavated anteriorly,
with a small denticular bump usually evident one
third the distance antero-posteriorly; having a
short but pronounced anterior siphonal trough;
with a flattened crescent-shaped area on base
adjacent to columella callous. Suture moderately
well impressed. Nuclear whorls light brown but
worn in all specimens examined. Operculum pau-
cispiral, dark brown and oval.
Details of animal and reproduction unknown.
‘XS 2 ae >| 5
Figs. 9,10. Smooth form of N. picta; Mokuoloe Id., Oahu,
Hawaii (USNM 346407, 7.2 x 4.7 mm.).
Figs. 11,12. Nodulose form of N. picta; same locality as
smooth form (USNM 346411, 8.5 « 5.6 mm.).
Figs. 13-18. Nodilittorina (Granulilittorina) cinerea (Pease).
Figs. 13,14. Lectotype of Littorina cinerea Pease [Mar-
quesas Islands] (ANSP 18811, 7.9 x 5.1 mm.).
Figs. 15,16. Moderately sculptured form, Atuona Bay,
Hivaoa Id., Marquesas (ANSP 155486, 8.9 x 5.7 mm.).
Figs. 17,18. Fatu Hiva, Marquesas (USNM 697101, 6.9 x
4.6 mm.).
Figs. 19,20. Holotype of Littorina iwakiana Nomura and
Hatai, from Tanagura Miocene, Japan (Saito Ho-on Kai
Museum, Sendai, Japan, Reg. no. 6895, 3.8 x 3.2 mm.).
___ 160°
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CAROLINE “Gas Tes
Plate 387. Geographical distribution of Nodilittorina (Granu-
lilittorina) exigua (Dunker),
[05-405 ]
502 Granulilittorina
Measurements (mm.)—
length width no. whorls locality
12.4 8.5 4+ Matsushima, Japan
11.2 7.6 4 Matsushima, Japan
10.8 Lio 4+ Komoi, Awaji, Japan
10.2 7.1 4 Matsushima, Japan
9.8 6.3 4 Peiyushan Id., China
9.1 6.2 4 Komoi, Awaji, Japan
8.8 6.3 4 Imaizumi, Japan
8.4 5.9 4 Shirahama, Japan
40 a 4 Hakodate, Japan
6.9 4.7 3+ Takami, Japan
Synonymy—
— Littorina granularis ‘Gray’ of Authors, non Gray, 1839;
Gray's holotype, BM(NH) 87.4.26.9 is the Eastern
Atlantic species, Nodilittorina miliaris (Quoy and
Gaimard),
1860 Litorina exigua Dunker, Malakozoologische Blatter
(1859), vol. 6, p. 226 (Japan); 1861, Dunker, Mollusca
Japonica, p. 13, pl. 2, fig. 3 [lectotype figure].
Joseph Rosewater
Littorinidae
Types—The type specimen of L. exigua
Dunker is probably in the Berlin Museum al-
though it has been impossible to locate it with
certainty during the present study. The specimen
figured by Dunker in the year following the
original description is here considered the lecto-
type (see pl. 386, figs. 1, 2).
Records—(All from USNM except where noted). CHINA:
Sidesaddle Island (Lu-hua_ shan), Chekiang Province;
Peiyushan Id.; Spider Island, Fukien Province; Big Wave
Bay, Hong Kong; Cape D’ Aguilar, Hong Kong. JAPAN: Hako-
date, Hokkaido; Saigo, Dogo Ids. Oki Group; Matsu-Shima;
Shirahama, Wakayama Province, Honshu; Toshima, Tanabe
Bay, Honshu; Awaji-shima; Kii, Honshu (AMS); Tokyo Bay
(ANSP); Imaizumi, Kagoshima Bay, Kyushu (USNM, ANSP);
Tosa Shikoku; Hachijo Island, 275 miles S. of Tokyo (both
ANSP).
[05-406]
November 30, 1970
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 503
Nodilittorina picta (Philippi, 1846)
(Pl. 386, figs. 7-12)
Range—Hawaiian Islands.
Remarks—The “Painted Littorine” Granulilit-
torina picta is closely related to G. millegrana, G.
exigua, and apparently also to G. cinerea Pease.
The four species all possess similar variation in
sculpture. The relationship with millegrana and
exigua was pointed out by Struhsaker (1968a)
who suggested that picta is the only one which
develops a smooth shell as well as a granulose one
[some populations of millegrana develop smooth
shells also; see pl. 386, fig. 7,8, 9, 10]. The possible
selective processes causing the extensive shell
variation in picta are well discussed by Struh-
saker (ibid.). It is probable that morphology of
most wild populations is under the positive con-
trol of natural selection there being few if any
mutations which survive for very long in nature
without some selective value. [The survival of so
many natural populations is coming under the
aegis of man, that the beneficial “weeding out”
effect of unsuitable elements by natural selection
may be considerably lessened. One wonders what
eventual effect this will have on all species in-
volved, including Homo sapiens.] It is, therefore,
most likely, as pointed out by Struhsaker that
each of the shell forms of picta, from highly
granulose to smooth, has survival value under
particular ecological conditions, i.e. smoother
forms survive best in wave-swept areas, and
sculptured forms occur in sheltered localities.
The morphological differences are apparently ge-
netically linked, but survival of the resultant
phenotype is under the control of environmental
selection.
The subspecies name L. picta marmorata Phi-
lippi has generally been used for one of the var-
iations of picta in Hawaii. The name marmorata
was not validly introduced by Philippi in connec-
tion with the original description of picta (see
synonymy) but was mentioned and figured in the
“Abbildungen” the following year with the add-
ed locality “provincia Ilocos borealis insulae
Lucon” [Philippines]. Both figure and locality
indicate that L. picta marmorata is a synonym of
G. millegrana. It is likely that Philippi, himself,
did not clearly discriminate between the popula-
tions of picta from Hawaii and some of the forms
of millegrana occurring in the Philippines. They
are indeed similar and this has given rise to
considerable confusion, wherein extra Hawaiian
Granulilittorina are called “picta.”’ As the mem-
bers of this subgenus all appear to be fairly
closely related it is perhaps a matter of personal
preference whether one considers them separate
species or geographic subspecies. In the case of
picta and its allies it is here considered less
confusing to consider them separate species.
In likening picta to neritoides, Philippi indica-
ted that he was describing a rather smooth little
shell, and the three syntypes of picta in the
British Museum (NH) reinforce this view. In my
experience, the name “marmorata”™ has been used
for the more highly sculptured forms, although
certainly the figure given by Philippi does not
show such a shell. It would appear that the local
interpretations of these names have come about
with usage. However, when the situation is ex-
amined closely it is found that picta is the only
valid name for this species; marmorata, as shown
above, not only is a synonym of another species
but is preoccupied by L. marmorata Pfeiffer,
1839 [= L. saxatilis Olivi]. Tinker’s (1952) sug-
gestion that picta is considered by some to be a
synonym of L. planaxis (eastern Pacific) is under-
standable. Certain populations especially of
young planaxis show similarities to picta in color
variation. They are, however, in distinct generic
groups, as well as being separate species (see List
of Taxa). The connection between L. planaxis and
picta probably dates from Tryon’s (1887) synony-
my.
Habitat—Shore rocks above high tide line (see
Struhsaker, 1968a).
Description—Shell reaching 12.9 mm. (about
0.5 inch) in length; rather conic to turbinate in
shape; average obesity about .64 (20 specimens
range from .59-.69); moderately thick in structure,
imperforate; surface often smooth, or only micro-
scopically spirally striate; but often bearing
raised, rather strong and granulose spiral cords on
spire and body whorls [granular sculpture most
similar to that in G. millegrana, but whorls
shouldered as in G. leucosticta and its subspe-
cies]. Axial sculpture consists of faint to rather
coarse, irregular lines of growth. External color
very variable: generally consisting of grayish to
yellowish white ground color “painted” with
dark-brown markings, a common pattern on body
whorl being the center of whorl dark, with area
above and below divided into light colored sec-
tions by wavy brown lines; The latter not at all
constant and brown painting may be minimized
or maximized; more highly sculptured shells ap-
pear darker overall. Aperture medium to dark
brown and may be brown and white mottled as is
[05-407]
504 Granulilittorina
exterior; with an often diffuse or interrupted
white band revolving inward from near anterior
junction of outer lip and columella; columella
usually light tan; inner edge of outer lip white or
with a few white spots. Base slightly flattened;
periphery occasionally with a weak to moderate
keel. Whorls 5-6, not very well-rounded, body
whorl distinctly flat-sided. Relative lengths of
spire and aperture about co-equal, one sometimes
exceeding the other apparently indiscriminately;
spire produced at an angle of from about 60-65°.
Aperture oval; outer lip moderately thick; inner
lip (columella) only moderately developed, witha
flattened crescent shaped area on base adjacent to
columella callus. Suture well impressed. Sculp-
ture varying from virtually smooth to markedly
spirally striate with coarse granulations. Nuclear
whorls light brown anteriorly, darker brown pos-
teriorly on each whorl, smooth and unsculptured;
in shells which will be granulosely sculptured,
this begins after about 21/2 nuclear whorls; in
shells which never develop coarse sculpture, only
low spiral striae begin after nuclear whorls. Oper-
culum paucispiral, brown, oval. Radula littori-
nid, 2-1-1-1-2, central tooth rather narrow.
Details of animal are from Whipple (1965): In
males base of penis is red as is testicular duct;
there is an enlargement at base of penis partially
separated from a penial gland containing an ac-
cessory flagellum; sperm groove runs along pos-
terior edge, deeply folded. Reproduction ovipa-
rous, producing a sculptured pelagic capsule
about 180m in diameter, with single egg about
75u in diameter. According to Struhsaker (1968b)
a swimming veliger hatches in about 3 days and
the larva settles and metamorphoses about 21
days after spawning.
Measurements (mm.)—
length width no. whorls locality
12.9 8.0 5+ Hawaiian Ids.
12.7 8.0 6 Waikiki, Oahu
11.8 7.9 5 Waikiki, Oahu
te 6.6 ay btwn Waipio and Pearl
City, Oahu
Joseph Rosewater
Littorinidae
10.9 6.8 5 Diamond Head, Oahu
10.0 6.8 5 Launiupoku, Maui
9.9 6.1 5 btwn Waipio and Pearl
City, Oahu
9.6 6.1 5 Paia Beach, Maui
9.1 5.9 6 Wainini, Kauai
Synonymy—
1846 Littorina picta Philippi, Proceedings of the Zoological
Society of London, for 1845, part 13, p. 139 (Hawaii);
lectotype in BM(NH) 1968324: 9.8 x 6.0 mm.; 1847,
as Litorina picta Philippi, Abbildungen und Besch-
reibungen Conchylien, vol. 2, Litorina p. 166 [not L.
picta marmorata, ibid., p. 167, pl. 3, fig 26 (from
Luzon, Philippines) = G. millegrana Philippi].
1857 Littorina picta Philippi, in Reeve, Conchologia Iconica,
vol. 10, Littorina pl. 15, figs 80 a, b, 81.
1887 Littorina planaxis ‘Nuttall’ Tryon, Manual of Concholo-
gy vol. 9, p. 248, pl. 44, fig 57 [only in part; includes
L. picta Philippi as synonym]; not L. planaxis ‘Nut-
tall’ Philippi, 1847, an eastern Pacific species.
Types—Three syntypes of Littorina picta Phi-
lippi were found in the British Museum (NH)
1968324. Of these, the one most nearly approxi-
mating the original measurements is here desig-
nated lectotype. Original measurements given by
Philippi were: “Altit. 4, diam. 3 lin.” which
yields the following millimeter measurements
based on a “German Line” equaling 2.18 mm.
(Rehder, 1945): length 8.7, width 6.6 mm. The
actual measurements of the designated lectotype
are 9.8 x 6.0 mm., fairly close to the original, and
here considered within the limits of a possible
measurement error. (see pl. 386, figs. 7, 8)
Records—HAWAIIAN ISLANDS: Laysan Island : La Pe-
rouse Rock, French Frigate Shoals; Nihoa Island (all BPBM).
KAUAI ISLAND: Nawiliwili (BPBM): Koloa (ANSP); Haena
(BPBM). OAHU: Kahuku Point (BPBM); Laie (ANSP, MCZ);
Kahana; Mokolii Island, Kaneohe Bay; Mokuoloe Island;
Moku Manu; Kailua; Manana (Rabbit Island) (all BPBM):
Diamond Head; Waikiki (both ANSP, MCZ, BPBM); Pearl
Harbor; Barber’s Point (both BPBM): off Puu Mailiilii (MCZ);:
E. of Waianae (ANSP); Makua (MCZ); Haleiwa (ANSP);
Waimea (BPBM). MOLOKAI: Pauwalu; Papohaku (both
BPBM); Moomomi (BPBM, ANSP); . MAUI: Lahaina (ANSP).
KAHOOLAWE ISLAND: (ANSP. HAWAII: Hilo (ANSP,
BPBM, MCZ); Kalapana; Waiahukini, Kau; Hoopuloa (all
BPBM); Refuge City, Honaunau Bay, Kona (ANSP); Keauhou
(BPBM); Kahaluu (ANSP); Kaulana, Kau; Tawai, Kalalaa, at
stream outlet (both BPBM).
[05-408 ]
November 30, 1970
Nodilittorina cinerea (Pease, 1869)
(Pl. 386, figs. 13-18)
Range—Marquesas Islands.
Remarks—N. (Granulilittorina) cinerea is
closely related to N. picta of Hawaii and shows
much of the same sort of variation from smooth to
granulose sculpture. A color pattern is also evi-
dent in occasional specimens of cinerea, but
never to the same degree as in picta. As in the case
of picta, cinerea is also quite closely related to the
other species of the subgenus Granulilittorina,
and some of its forms are very close to N. mil-
legrana. Major distinguishing features are (1) the
gray color as indicated by Pease’s name “‘ci-
nerea’, which never reaches the marbled colora-
tion of picta, (2) the whorls which may become
relatively broadly shouldered, and (3) the degree
of sculpture, ranging from almost smooth to fairly
granulose spiral cords. According to available
records, this species is endemic in the Marquesas
Islands.
Habitat—In splash zone, above high tide line,
in cracks in lava, crawling on rock surface at night
(personal communication, H. A. Rehder; from
observations made at Nuku Hiva, Marquesas Is-
lands, 1967).
Description—Shell reaching 9.0 mm. (about /3
inch) in length; turbinate to shouldered-turbinate
in shape; average obesity about .65 (20 specimens
range from .58-.71); only moderately thick in
structure; imperforate; sculpture of low to rather
distinct spiral cords; cords smooth or granulose;
whorls varying from only slightly to rather dis-
tinctly shouldered; axial sculpture consisting of
faint to coarse growth lines and furrows; occa-
sionally specimens have a deep, ragged axial
mark of growth interruption. External ground
color somewhat variable from grayish white to
distinct bluish gray, the latter especially in the
young; occasional patterning consists of irregular
darker gray to brown zigzag lines and flammules.
Aperture medium reddish brown to dark brown
with a white band revolving inward from near
anterior junction of outer lip and columella; colu-
mella usually colored about as aperture or slightly
lighter; inner edge of outer lip white. Base some-
what flattened; periphery not keeled, whorls
shouldered for about #/s the distance anterior to
suture. Whorls about 4-6, not too well rounded
but rather tending to be straight-sided. Relative
INDO-PACIFIC MOLLUSCA, vol. 2, no. 11
Nodilittorina 505
lengths of spire and aperture about coequal or
spire slightly shorter. Spire produced at an angle
of from about 60-67°. Aperture widely oval; outer
lip moderately thick; inner lip (columella) moder-
ately strong, with an often well depressed cres-
cent shaped area on base adjacent to columella
callous; the depression rarely approaching an
umbilical chink. Suture well impressed. Sculp-
ture varying from specimens with hardly raised
spiral cords to those exhibiting distinct well
raised cords with granulations on all whorls.
Nuclear whorls light brown anteriorly, dark
brown posteriorly smooth and unsculptured; nu-
cleus about 2 whorls in extent; first postnuclear
whorl smooth, subsequent whorls with spiral
sculpture; overall microscopic spiral sculpture
consisting of fine wavy spiral threads. Operculum
paucispiral light brown, oval. Radula littorinid,
2-1-1-1-2, central tooth rather narrow.
Animal similar to other Nodilittorina. Nothing
is known concerning reproduction and _ larval
development; probably similar to G. picta.
Measurements (mm.)—
length width no. whorls locality
9.0 5.8 5+ Hiva Oa, Marquesas
8.7 5.3 4+ Hiva Oa, Marquesas
8.5 5.6 5+ Hiva Oa, Marquesas
8.2 4.8 6+ Hiva Oa, Marquesas
8.1 5.2 4+ Hiva Oa, Marquesas
7.9 5.1 6 Lectotype, Marquesas
13 bd. 4+ Fatu Hiva, Marquesas
7.0 AT 3+ Fatu Hiva, Marquesas
6.8 4.6 4+ Fatu Hiva, Marquesas
6.4 4.3 4+ paralectotype,
Marquesas
Synonymy—
1869 Littorina cinerea Pease, American Journal of Concholo-
gy, vol. 5, part 2, p. 78, pl. 8, fig. 14 (Marquesas
Islands); lectotype ANSP 18811: 7.9 x 5.1 mm.
Types—tThere are three syntypes of Littorina
cinerea Pease in the ANSP. Of these, one, most
resembling the figure associated with the original
description, is here designated lectotype: ANSP
18811 (pl. 386, figs. 13, 14). Its measurements:
length 8.1, width 5.2, are fairly close to the
originals given by Pease: 7 x 5.5 mm. The lecto-
type and two paralectotypes fairly well fulfill
Pease’s descriptive comments: “... . transversely
granosely ridged or smooth....
whitish, apex blackish ..... °
Records—MARQUESAS ISLANDS: Nuku Hiva (ANSP;
USNM);: Ua Pou; Tahuata (both USNM):; Ua Huka (ANSP);
Hiva Oa (ANSP, USNM); Fatu Hiva (USNM).
cinereous or
[05-409]
506 Granulilittorina
Joseph Rosewater
Littorinidae
Nodilittorina iwakiana
(Nomura and Hatai, 1936)
(Pl. 386, figs. 19, 20)
Range—Miocene, Japan.
Remarks—The tiny species described by No-
mura and Hatai appears to show closest affinity
with the members of the subgenus Granulilit-
torina. Details of sculpture given by the authors
suggest that this fossil may be related to G.
millegrana Philippi.
Synonymy—
1936 Littorina iwakiana Nomura and Hatai, Saito Ho-on Kai
Museum Research Bulletin, no. 10, p. 144, pl. 16, figs
16a, b. (Tanagura Miocene, Okada, Japan); unique
holotype in Saito Ho-on Kai Museum, Sendae, Japan,
Reg. No. 6895; 3.8 x 3.2 mm.
Published by
THE DEPARTMENT OF MOLLUSKS
Delaware Museum of Natural History
Greenville, Delaware
19807, U.S.A.
[05-410]
January 15, 1972
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
Tectariinae 507
WILL IAM H. |}
SECTIONAI
DIVISIO
4
THE FAMILY LITTORINIDAE IN THE INDO-PACIFIC
Part Il. The Subfamilies Tectariinae and Echinininae
by JOSEPH ROSEWATER
Division of Mollusks
National Museum of Natural History
Washington, D.C. 20560 U.S.A.
Abstract
The classification of tropical Indo-Pacific
Tectariinae and Echinininae (Mollusca: Meso-
gastropoda: Littorinidae) has been revised. Seven
Recent and three Tertiary fossil species belonging
to three generic or subgeneric groups are rede-
scribed and figured. Complete systematic synon-
ymies are given, together with discussions of
relationships, biology and zoogeography. A list is
given of world-wide members of Tectariinae and
Echinininae.
Introduction
In Part I of Indo-Pacific Littorinidae the spe-
cies belonging to the subfamily Littorininae were
reviewed by Rosewater (1970). The present sec-
tion covers the subfamilies Tectariinae and Ech-
inininae. As yet not covered are such groups as
Bembicium and its allies and Cremnoconchus,
although they both occur in the Indo-Pacific re-
gion. They are here considered to constitute at
least subfamily groups, if not separate families
(see Bibliography in Rosewater, ibid., p. 427:
Anderson, D. T., 1960, and also Anderson, H.,
1958; also see Prashad, 1925 and Kesteven, 1903).
The many Antarctic littorinids were excluded
from Part I of this study since actually they are
out of the geographic area of the tropical Indo-
Pacific (see Powell, 1951 and Dell, 1964).
The Littorininae, Tectariinae and Echinininae
differ in habitat preference and in morphology in
a number of aspects. Echinininae tend generally
to live rather high on the shore. Tectariinae are
next and Littorininae usually are nearest to the
sea, although considerable variation exists, the
genus Nodilittorina having some high-living spe-
cies. The differentiation of the three subfamilies
also is based on the following morphological
grounds. In Littorininae the radula is of the gen-
eralized littorinid type with some narrowing of
the central tooth in Nodilittorina; opercula are
paucispiral; penises are adorned with a few peni-
al glands but otherwise are not particularly com-
plicated.
In the Tectariinae the lateral radula tooth is
partitioned and appears thickened and differently
oriented than in either Littorininae or Echin-
ininae; the opercula are mesospiral (see below);
penises are abundantly supplied with glands and
are also papillose on non-glandular surfaces. The
shells in both Littorininae and Tectariinae usual-
ly are imperforate although the subgenus Cen-
chritis is an exception in the latter subfamily.
In Echinininae the radula is not unusual, the
subgenus Tectininus excepted, where reduction
has taken place; opercula are multispiral; shells
are umbilicate, usually; penises are supplied with
a number of penial glands near the base. In the
possession of an umbilicus Cenchritis could be
considered to form a bridge between Tectarius
and Echininus. In general, however, Tectarius
appears to be more closely related to Littorina in
its conservative characters, while Echininus may
be considered a specialized group perhaps evolv-
ing toward a land environment. These three
subfamily groups provide a convenient and ap-
parently natural framework upon which to ar-
range the contained genera and species (see illus-
[05-441]
508 Joseph Rosewater Littorinidae
Plate 388. Subfamilies Tectariinae and Echinininae
(explanation on opposite page; all figures about natural size)
[05-442]
January 15, 1972
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
Tectariinae 509
trations of these morphological differences in
Rosewater, 1970, and in this paper).
Most Tectarius s.s.and Echininus s.s. are living
today only in the East Indian area. With the
exception of Tectarius grandinatus whose range
extends eastward to Polynesia, all other species
are inhabitants of the raised, weathered coral reef
shorelines found in the Western Pacific Arc (per-
sonal observations, 1970). It can only be assumed
that this niche provides the requirements essen-
tial for the existence of these species‘as they are to
be found nowhere else.
The fossil record provides very few clues to the
origin of these groups. There are only three Terti-
ary fossil species described from the Indo-Pacific
and these probably belong in three separate ge-
neric taxa. The oldest of these, T. songoense
Martin, from the Upper Eocene of Java, probably
represents nearly the earliest appearance of Tec-
tarius. As mentioned in Part I, littorinid fossils
are exceedingly difficult to separate from Trochi-
dae and Turbinidae, and this is no less true for
Tectarius and Echininus. Probably, however,
these groups made their appearance in the early
Tertiary within the region where they have devel-
oped, and with the exceptions of Echininus (Tec-
tininus) nodulosus and Tectarius (Cenchritis)
muricatus, both of the western Atlantic, they have
remained there.
Opercula
The opercula of Indo-Pacific Littorinidae re-
quire special comment (see pl. 389). All are made
up of conchiolin, and those of members of the
subfamily Littorininae, including Littorina, its
subgenera, and Nodilittorina are usually pauci-
spiral and rather oval in shape with the nucleus at
the side and nearer one end (oligogyrous spiral
type of Fretter, et al., 1962, pp. 79,80). In the
Echinininae, the basic plan of the operculum
differs from that of other littorines. It is the type
Plate 389. Opercula,of Littorinidae and Trochidae.
Fig. A. Paucispiral operculum of Littorina (Littorinopsis)
scabra (Linné) from Mokuoloe Id., Kaneohe Bay, Oahu
(USNM 339388).
Fig. B. Mesospiral operculum of Tectarius rusticus (Philip-
pi) from Troughton Chain, northern Western Australia
(WAM 1787-69);
Fig. C. Multispiral operculum of Echininus cumingi (Phi-
lippi) from near Davao City, Mindanao, Philippines
(WAM 1566-70);
Fig. D. Multispiral operculum of Trochus niloticus Linne,
from Makuluva, Viti Levu, Fiji (USNM 531827).
Lines under each figure represent 5 mm.; stippled areas
are thickened and dark-brown in color; non-stippled areas
are ight horn color and transparent.
of operculum which is called in other groups,
such as Trochidae, a multispiral operculum
(polygyrous spiral type of Fretter, et al., ibid.)
although not so extreme as that figured by Fetter
(ibid., p. 80, fig. 43A; also see our pl. 389, fig. D.).
The operculum is circular in outline and moder-
ate to small in size. The nucleus is decidedly
central in location and growth proceeds outward
from the center in multiple, fairly evenly spaced
gyrations (pl. 389, fig. D). The operculum in Tec-
tarininae (fig. B) is intermediate in form between
Explanation to plate 388 (opposite page)
Figs. 1,2. Tectarius grandinatus (Gmelin) from Palmerston
Atoll, Cook Islands (USNM 685165)
Figs. 3,4. Tectarius tectumpersicum (Linné). Fig. 3, from
Stirling Isle, Treasury Ids., Solomon Islands (USNM
600370); Fig. 4, from “East Indies” (USNM 131450).
Figs. 5-7. Tectarius pagodus (Linné). Fig. 5, from “East
Indies” (USNM_ 18966); Fig. 6, from the Philippines
(USNM): Fig. 7, a young specimen from Polillo, Philip-
pines (USNM 311141).
Figs. 8,9. Tectarius rusticus (Philippi) from Buccaneer
Archipelago, Western Australia (USNM 684713).
Figs. 10,11 Tectarius rugosus (Wood). Fig. 10, from Pacific
(USNM 304587); Fig. 11, from Davao Bay, Mindanao,
Philippines (USNM 654034).
Figs. 12,13. Echininus cumingi cumingi (Philippi) from
Hervey Ids., Cook Ids. (USNM 42452).
Figs. 14,15. Echininus cumingi spinulosus (Philippi) from
Kadena Circle, Okinawa, Ryukyu Ids. (USNM 664658).
Figs. 16,17. Tectarius (Cenchritis) muricatus (Linne), from
Boca de Camarioca, Matanzas, Cuba (USNM 599944).
Figs. 18-20. Echininus (Tectininus) nodulosus (Pfeiffer).
Fig. 18, from Hog Island, Bahamas (USNM 603911);
Figs. 19, 20, from Mujeres Harbor, Quintana Roo, Mexico
(USNM 662308).
[05-443]
510
Joseph Rosewater
Littorinidae
Littorininae and Echinininae, being large and
rather rounded in outline, with a slightly acentric
nucleus and having a number of gyrations more
than the paucispiral type but less in number than
the multispiral type. The Tectariine opercular
type is here termed the mesospiral or mesogyrous
spiral type. The presence of the three opercular
types in Littorinidae may be considered to have
evolutionary significance, and possibly is related
to selection for a better aperture sealing mecha-
nism in animals which have considerable vertical
distribution on the shore line: in order, proceed-
ing from low toward higher shore habitats—
Littorininae, Tectariinae, Echinininae.
Reproduction
To my knowledge nothing is known concern-
ing reproduction in either Tectariinae or Echin-
ininae with the exception of Tectarius (Cen-
chritis) muricatus (Linné) which produces a pe-
lagic capsule (see Lebour, 1945, and Lewis, 1960,
references in Rosewater, 1970 p. 05-276). Field
and laboratory studies are needed to discover
details of the life histories of the remaining spe-
cies. However it is likely that most of these snails
also produce eggs encased in pelagic capsules
which undergo development in the sea. Abbott
(1954) noted that Lebour (ibid.) stated that some
of the Bermuda littorinids that live above high
tide line migrate to the water to spawn. It is
suspected that this also is the case with many of
the Indo-Pacific species.
Acknowledgments
The persons and institutions acknowledged in
Part I of this study (see Rosewater, Indo-Pacific
Mollusca, vol. 2, no. 11, p. 425) also are thanked
here. In addition, I acknowledge the following
for their help in making possible the examination
in the field of most of the species of Tectarius and
Echininus during the National Geographic Socie-
tvy—Mariel King Memorial Expedition to the Mo-
luecas Islands, Indonesia, May to July 1970: the
late Mariel King, Mrs. Grace King, T. H. Richert,
C. Beal, C. M. Burgess, B. R. Wilson, and the
National Geographic Society. The Government of
Indonesia graciously provided clearance for the
vessel Pele to work in the Moluccas Islands. Mr.
Kasim Moosa and Mr. Sukarno, both of the Insti-
tute for Marine Research, Djakarta, accompanied
the expedition and provided assistance of many
kinds.
List of Recognized Taxa
Below is a list of the Tertiary fossil and Recent
species herein recognized as belonging in the
subfamilies Tectariinae and Echinininae. The
few fossil taxa are preceded by a dagger [f].
Family Littorinidae Gray, 1840
Subfamily Tectariinae, new subfamily
GENUS Tectarius Valenciennes, [1832]
Subgenus Tectarius Valenciennes, [1832]
rugosus (Wood, 1828). Type. Recent, wes-
tern Pacific
grandinatus (Gmelin, 1791). Recent, Pacific
islands
pagodus (Linné, 1758). Recent, western Pa-
cific
tectumpersicum (Linné, 1758). Recent, wes-
tern Pacific
rusticus (Philippi, 1846). Recent, northern
Australia
+ songoense (K. Martin, 1931). Eocene, Java.
Subgenus
+Subditotectarius Ladd, 1966
trehderi Ladd, 1966. Type. Miocene, Mar-
shall Islands.
Subgenus Cenchritis von Martens, 1900
muricatus (Linne, 1758). Type.
tropical western Atlantic.
Recent,
Subfamily Echinininae, new subfamily
GENUS Echininus Clench and Abbott, 1942
Subgenus Echininus Clench and Abbott, 1942
cumingi cumingi (Philippi, 1846). Type. Re-
cent, western Pacific
cumingi spinulosus (Philippi, 1847). Recent,
western Pacific
+ adelaidensis (Cotton, 1947). Pliocene, South
Australia,
Subgenus Tectininus Clench and Abbott, 1942
nodulosus (Pfeiffer, 1839). Type. Recent,
tropical western Atlantic.
[05-444]
January 15, 1972
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Rosewater, J. 1970. The Family Littorinidae in the Indo-
Pacific. Part I. The Subfamily Littorininae. Indo-Pacific
Mollusca, vol. 2, no. 11, pp. 417-506, 64 pls.
Rumphius, G. E. 1705. D’Amboinsche Rariteitkamer. Am-
sterdam, 340 pp., 60 pls.
Sherborn, C. D. and B. B. Woodward. 1901. Bibliographical
Notes. XXVIII. The Dates of Humboldt and Bonpland’s
“Voyage”. Journal of Botany, June, pp. 1-4.
Troschel, F. H. 1856-1863. Das Gebiss der Schnecken, vol. 1,
Berlin, pp. vii + 252, pl. 1-20.
Watson, R. B. 1886. Report on the Scientific Results of the
H.M.S. Challenger, vol. 15, part 42, p. 576.
Wenz, W. 1938. Handbuch der Paliozoologie, vol. 6, part 1,
lfg. 3, pp. 241-480, figs. 472-1235.
Wimmer, August. 1880. Sitzungsberichte der Mathematisch-
naturwissenschaftlichen Classe der Kaiserlichen Akademie
der Wissenschaften, Wien, I. Abth. vol. 80, pp. 496, 514.
[05-447]
512 Joseph Rosewater Littorinidae
Key to the Tectariinae and Echinininae
The following key is to the genera and sub- only, since Western Atlantic taxa are not treated in
genera of these two subfamilies. It is based upon detail in the present paper. For a key to the
the shell and externally observable characters. Littorininae, see vol. 2, no. 11, p. 430 [p. 05-278].
Page numbers are given for Indo-Pacific groups
aS etl (iri WCAC ye cus. ea dd a Rhaclea eatacte. Shin al ites dachass lee yh Bh cee een eee 2
Lb: Shell Wor aii TCAte be «scm ted 6 ph ated ate aime we cheer sek De Dh eas Se es ee 3
2a Shell spinose, with partially open spines, shell about as wide as high, operculum
maul tigate 166C4pl G00) .< ¢ hae 4 Gnd 4.44 24 4A Ae es Sas Echininus p. 526
2b Shell not spinose, nodulose, higher than wide, operculum not multispiral ... Cenchritis
3a Shell spinose, operculum mesospiral. Tectarius ......0. 5 ccc enna prods
oe Shell moculoce: (ose) 3c as swe whee one oe Hab ie 4 PGloe otis Subditotectarius p. 524
3c Shell moderately spinose, operculum multispiral................0.05. Tectininus
Plate 390. Type-species of Genera and Subgenera of Tectari- Miocene, Marshall Islands (Holotype, USNM 648342; 2.8
inae (Figs. 1-9) and Echinininae (Figs. 10-15) illustrating x 2.4 mm.).
sculpture, arrangement of spines, and presence or absence of Figs. 7-9. Tectarius (Cenchritis) muricatus (Linné);
umbilici. Matanzas, Cuba (USNM 599944; 26.1 «x 17.9 mm.).
Figs. 10-12. Echininus (Echininus) cumingi (Philippi);
Figs. 1-3. Tectarius (Tectarius) rugosus (Wood); Davao Bay, Cook Islands (USNM 42452a; 17.2 « 17.5 mm.).
Mindanao, Philippines (USNM 654034; 23.1 x 17.8 mm.). Figs. 13-15. Echininus (Tectininus) nodulosus (Pfeiffer);
Figs. 4-6. Tectarius (Subditotectarius) rehderi Ladd; early Cozumel Id., Mexico (USNM 662806; 14.4 x 12.2 mm.).
[05-448]
January 15, 1972
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
Tectarius 513
Subfamily Tectariinae, new subfamily
Genus Tectarius Valenciennes, [1832]
Type: Tectarius rugosus (Wood, 1828)
The genus Tectarius sensu lato includes the
Tectarius, whose type-
species T. rugosus Wood (= T. papillosus ‘La-
marck’ of authors) has one of the more conserva-
tively sculptured shells of the group. Tectarius
sensu stricto is a wholly Indo-Pacific group, hav-
ing its present population center in the Western
Pacific Arc. The monotypic subgenus Cenchritis
von Martens contains only T. (C.) muricatus
(Linné), of the tropical western Atlantic (see pl.
388, figs. 16, 17). Subditotectarius Ladd, 1966, is
monotypic for the fossil T. (S.) rehderi Ladd, of
the Miocene of the Marshall Islands. Only spe-
cies belonging to Tectarius s.s. and Subditotec-
tarius will be considered here.
Tectarius appears more closely related to Lit-
torina than to Echininus because of greater simi-
larities in morphology, general shell characters,
the absence of a truly multispiral operculum, a
broader, less modified central radula tooth, and
the usual lack of an openly umbilicate shell.
nominate subgenus,
Subgenus Tectarius sensu stricto
Moderately large, pyramidal to turbinate,
non-umbilicate littorinids with from rather
strongly spinose to nodulose or papillose shells;
generally living at or above high tide line. Radula
littorinoid, the central tooth somewhat narrowed,
the lateral tooth with an embayment and usually
developing a medial vertical ridge or partition. In
males the penis is large and well-supplied with
glands along most of its lateral edge, the remain-
der papillose, and with an open but deeply folded
seminal duct. Operculum rounded, mesospiral
(see Opercula in Introduction). Aperture plicate
within; with a columellar swelling or tooth.
Synonymy—
1798 Cidaris Roding, Museum Boltenianum, part 2, p. 84;
type-species by subsequent designation, Herrmann-
sen, 1847: Trochus pagodus Linné; non Cidaris
Leske, 1778, nor Swainson, 1840.
[1832] Tectarius Valenciennes, Coquilles, in Humboldt and
Bonpland. Voyage aux régions équinoxiales du
Noveau Continent, Observations de Zoologie, vol. 2,
p. 271; type-species by subsequent designation
Clench and Abbott, 1942: Trochus coronatus Valenci-
ennes [= Tectarius rugosus (Wood]. ICZN Opinion
871.
1839 Pagodus Gray, in Molluscous Animals: The Zoology of
Captain Beechey’s Voyage, p. 141; type-species by
Monotypy and by absolute tautonymy, Monodonta
pagodus Lamarck [= Tectarius pagodus (Linné].
1840 Pagodella Swainson, A Treatise on Malacology, pp. 207,
219, 221 [refers to Pagodella echinata, nomen nudum|
351; refers to P. major Martini - Chemnitz, pl. 163,
figs 1541, 1542 [= Tectarius pagodus (Linné)] and to
T. tectumpersicum ibid., fig. 1543, 1544; type-species
here designated: Tectarius pagodus (Linné).
1840 Echinella Swainson, ibid., pp. 207, 221, 352; refers to E.
granulata Swainson [nomen nudum] and to E. coro-
naria, Tableau Encyclopédique et Méthodique, pl.
447, fig 6 [=Mondonta coronaria Lamarck = Tectar-
ius grandinatus (Gmelin)]; type-species by monotypy,
Tectarius grandinatus (Gmelin) [also see Clench and
Abbott, 1942]; not Echinella Bory St. Vincent, 1824.
1846 Fectaria Philippi, Abbildungen und Beschreibungen
Conchylien, Vol. 2, Litorina, p. 139; used in combina-
tion Fectaria pagodus; error for Tectarius Valencien-
nes,
1858 Hamus ‘Klein’ H. & A. Adams, The Genera of Recent
Mollusca, vol. 2, p. 656, refers to H. and A. Adams,
1854, vol. 1, p. 315; type-species here designated,
Hamus pagodus (Linné) [=Tectarius pagodus
(Linné)]; not Hamus ‘Klein? R. B. Watson, 1886
[=Trochidae].
1899 Echinellopsis Rovereto, Atti della Societa Ligustica di
Scienze naturali e geografiche, vol. 10, p. 109; new
name for Echinella Swainson, 1840, not Bory St.
Vincent, 1824.
Nomenclature—Due to similarities between the
shells of Tectarius and some of the Trochidae,
there has been a tendency for some of the former
to be classified with the latter. This problem was
discussed by Keen (1966) who recommended that
the International Commission on Zoological
Nomenclature validate Tectarius with the type-
species Tectarius coronatus Valenciennes,
[1832], i.e., in its accustomed sense. Her petition
was granted in I.C.Z.N. Opinion 871 (Melville
and China, 1969). It was assumed in this Opinion
that the type-species of Tectarius, T. coronatus
Valenciennes, is a synonym of T. grandinatus
Gmelin. However, an examination of the type-
specimen of coronatus in the Paris Museum
shows it to be in actuality T. rugosus Wood,
which usually has been erroneously referred to as
T. papillosus Lamarck.
Another name which sometimes has been as-
sociated with Tectarius that has an exceedingly
long and complicated history is the genus Ha-
mus. It was mentioned originally by Klein (1753)
where its use was of course pre-linnaean. Bru-
guiére (1792) gave a brief description, referring to
Klein, but listed no species. Deshayes (1830)
[05-449]
514 Joseph Rosewater
Littorinidae
declared it “‘a forgotten genus’, indicating that he
considered it unrecognizable. Mérch (1852)listed
it, this time in the synonymy of Littorina Férus-
sac, an invalid introduction (I.C.Z.N., Art. 11(d)).
The first valid use of Hamus was not until H. & A.
Adams (1858) used it as a senior synonym for
Tectarius (see synonymy). It was later used by
Wimmer (1880) and Watson (1886), the last being
a taxon of Trochidae. I have designated as type-
species of Hamus H. and A. Adams, 1858, H.
pagodus (Linné) and consider this genus to be an
absolute synonym of Tectarius Valenciennes.
The use of square brackets surrounding the
date for Tectarius Valenciennes, [1832] is recom-
mended by the International Code of Zoological
Nomenclature in cases where the date of publica-
tion of a name has been determined on the basis
of external evidence (I.C.Z.N. Recommendation
22.A(3); also see Sherborn and Woodward, 1901;
Keen, 1966; and Opinion 871).
Tectarius rugosus (Wood, 1828)
(Pl. 388, figs. 10, 11)
Range—Philippines and Indonesia.
Remarks—The shells of well prepared and
cleaned specimens of Tectarius rugosus tend to
be quite colorful for Littorinidae, with the
orange-pink coloration of the last two whorls
contrasting with a purplish brown subsutural
band. These colors do not show well in all speci-
mens, however, and are not very visible in un-
cleaned specimens. The three’ large, non-
umbilicate common species of the southwest Pa-
cific may be distinguished by the number of
major spiral rows of spines on the last whorl: 2 in
pagodus; 3 in tectumpersicum; and 4 in rugosus.
The closely-spaced stubby spines of rugosus also
separate it from the other two. These characteris-
tics do not of course help to distinguish it from T.
grandinatus, but other characters and the Polyne-
sian endemicity of the latter are helpful in this
case (see Remarks under grandinatus).
Habitat—Shore rocks and limestone cliffs 1-2
meters above high tide line (personal observa-
tions, Davao, Philippines, 1970).
Description—Shell reaching 39.7 mm (about
1?/2 inches) in length, broadly conical in shape,
average obesity about .76 (51 specimens range
from .68-.82); mature specimens moderately
heavily constructed, imperforate, and sculptured
on most postnuclear whorls with four, fairly
closely-spaced rows of stubby, rounded, often
slightly upturned spines. External color generally
yellowish white on early whorls, becoming pink-
ish orange on penultimate and body whorls; area
of most posterior (subsutural) row of spines usual-
ly a contrasting purplish brown, and the same
dark color may appear in lines and dashes inside
outer lip of aperture; aperture tinted lighter pink-
ish orange. Base flattened, sculptured spirally
with nodulose cords, a larger separate row just
below periphery of body whorl. Whorls 6-8, flat-
sided excepting spines. Length of spire usually
greater than half the length of shell. Spire convex,
produced at an angle of from about 60-67°. Aper-
ture rounded-squarish; outer lip thickly produced
in mature individuals, strongly plicate within;
plicae not reaching edge of aperture; outer lip
tapering to a thin, crenulate edge; inner lip
smooth posteriorly, often stained a deeper orange
than rest of aperture, forming a tooth-like bulge
anteriorly, near junction with outer lip near base
of columella. Suture obscured by anteriormost
row of spines of preceding whorl. Primary sculp-
tural feature is the four spiral rows of spines.
Spines not particularly aligned axially, although
anteriormost 2 rows more so than others; from
17-23 spines per row on body whorl; bases of
anteriormost 3 rows of spines joined by low spiral
carinae. Posteriormost-but-one (3rd) row of
Plate 391. Tectarius rugosus (Wood, 1828).
Figs. 1,2. Turbo rugosus Wood, lectotype, BM(NH)
1968370, 28.4 x 22.3 mm.
Figs. 3,4. Tectarius coronatus Valenciennes, Holotype,
MHNP (“‘Acapulco” [Luzon, Philippines]) 32.9 x 25.7
mim.
[05-450]
January 15, 1972
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
Tectarius 515
spines protrudes farthest on spire whorls, but this
distinction largely lost on more mature whorls;
posteriormost (4th) row of spines obscured by
subsutural purplish brown color band especially
on penultimate and body whorls. Secondary
spiral sculpture, between each row of spines, con-
sisting of raised cords, and overall spiral sculp-
ture of microscopic threads. Axial sculpture con-
sists of irregular flaky lines of growth. Operculum
moderate in size, circular, an average one measur-
ing about 7 mm. in diameter, thin, light-brown
with a dark-brown center, paucispiral, nucleus
about central. Periostracum not evident. Nuclear
whorls at least partially decollate in all specimens
examined, about 2, smooth, grayish white, first
postnuclear whorls showing early signs of spiral
striae and becoming nodulose. Radula littorinoid,
9-1-1-1-2; lateral tooth with a vertical partition
and an embayment typical of Littorinidae. Ani-
mal moderately large, littorinoid; penis large,
muscular and apparently highly extensible; semi-
nal groove in deep fold running along medial
edge to tip; tip vermiform, covered with papillae;
lateral edge of penis supplied with large number
of glands not extending onto vermiform tip. Re-
productive data and life history unknown.
Measurements (mm) (all Philippines)—
length width No.
whorls
locality
39.7 27.6 7+ Cadao Id., Naro Bay, Masbate
35 25,2 7+ Cadao Id., Naro Bay, Masbate
30.1 22.4 7+ San Miguel Bay, Ticao
95.9 91.2 8 Batag Id., Samar
23.9 16.3 T+ Bongao Channal, SW Sanga Sanga
Id., Sulu Archipelago
21.0 15:3. 7+ Borongan, E side Samar
io ikea ee Papahag Id., Tawi Tawi Group
15.0 12.0 6+ Borongan Village, E side Samar
13:5 10.8 ate Papahag Id., Tawi Tawi Group
12.6 9.5 6+ Papahag Id., Tawi Tawi
La? 8.4 6+ Papahag Id., Tawi Tawi
8.4 6.8 5+ Papahag Id., Tawi Tawi
Synonymy—
Monodonta papillosa of authors, not M. papillosa
Lamarck, 1822 [=Tectarius tectumpersicum (Linne,
1758) ].
1828 Trochus rugosus Wood, Supplement to the Index Tes-
taceologicus or a Catalogue of Shells, British and
Foreign, pl. 5, Trochus, fig. 7 (no locality given;
Mindanao, Philippines, here selected); lectotvpe in
BM(NH) 1968370, length 28.4 mm, width ( ca. ) 22.3;
not Litorina rugosa Menke, 1843 [= Nodilittorina aus-
tralis (Gray, 1826)].
1832 Tectarius coronatus Valenciennes in Humboldt and
Bonpland, Voyage aux régions equinoxiales du
Noveau Continent, vol. 2, Coquilles, p. 271 (Acapulco
[in error] locality here corrected to Luzon, Philip-
pines); Holotype in MHNP.
1846 Litorina papillosa elegans Philippi, Abbildungen und
Beschreibungen Conchylien, vol. 2, p. 140, Litorina,
pl. 2, figs. 5, 7 (precise locality not given); figured
specimens from Cuming Collection BM(NH) [not
seen during 1968 visit] and Saul Collection, Cam-
bridge Museum.
1846 Litorina papillosa quadriseriata Philippi, ibid., p. 140,
Litorina, pl. 2, fig. 2 (Zanzibar [in error] locality here
corrected to Luzon, Philippines); type-specimen may
be in BM(NH) [not seen during 1968 visit]; refers to
“Trochus rugosus Wood Suppl. t.5, £. 7”.
Types—Although the name T. papillosus has
been applied to this species (see Kaicher, 1956)
the type-specimen of that species in the Geneva
Museum is unquestionably T. tectumpersicum
Linné, and papillosus is, therefore an absolute
synonym of tectumpersicum (q.v.). It also has
been referred to as Echinellopsis grandinatus
(Habe, 1961, p. 20; 1964, p. 28, both pl. 9, fig. 30)
which is an error of nomenclature for T. rugosus.
The first available name is Trochus rugosus
Wood, 1828, the lectotype of which is in the
BM(NH) 1968370. The holotype of T. coronatus
Valenciennes is in the Paris Museum. The figured
specimens of Philippi’s elegans and quadriseriata
may be in the BM(NH) and/or the Cambridge
Museum. They were not discovered by me at the
BM and may be lost. Philippi’s figures are quite
adequate for the interpretation of the species and
may be considered as representative of the lecto-
types: elegans, pl. 2, fig. 7; quadriseriata, pl. 2,
fig. 2.
1oo° 120° \4o° 16¢
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) MARIANA
is
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NICOBAR |; cack e
16 % Tt te Pon
CEYLON ise, |e
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COCO6 - KEELING
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100 120° 140° 160
Plate 392. Geographic distribution of Tectarius rugosus
(Wood) in the Philippines and Indonesia.
[05-451]
516 Joseph Rosewater
Records—PHILIPPINES: Port Galera, Mindoro; San
Miguel Bay, Ticao Id.; Cadao Id., Naro Bay, Masbate (all
USNM); Borongan Village (USNM, Del. Mus. N.H., ANSP,
MCZ); Batag Id., both Samar (USNM); Samal Id., Davao Bay
(MCZ, USNM, WAM); Zamboanga, both Mindanao (ANSP,
Del. Mus. N.H.); Jolo Id. (MCZ); Tabawan Id. (ANSP);
Papahag Id. (USNM); Bongao Channel, SW Sanga Sanga Id.,
all Sulu Archipelago (ANSP). INDONESIA: Bula Buka Id.
Gulf of Tomini, Celebes (USNM).
Tectarius grandinatus (Gmelin, 1791)
(Pl. 388, figs. 1,2; pls. 393-395)
Range—The Cook, Society, Tuamotu and
Gambier Islands, southeastern Polynesia.
Remarks—Tectarius grandinatus, an inhabitant
of southeastern Polynesia, apparently is ge-
ographically isolated from the several other mem-
bers of the genus Tectarius in the Indo-Pacific,
the others being found in the East Indies. It is
quite distinct, morphologically, from pagodus
and tectumpersicum, but, interestingly, is very
close in appearance to T. rugosus in general
matters of sculpture and external anatomy of the
animal. Superficially, grandinatus might be
thought more closely related to Cenchritis muri-
catus (L.) of the western Atlantic. However, the
radula, penial anatomy (Abbott, 1954) and a close
examination of shell sculpture and _ structure
causes me to reject that theory: grandinatus has
not been observed to be umbilicate, while muri-
catus sometimes is; the aperture of muricatus is
never thickened and plicate as it is in grandinat-
us; the operculum of grandinatus is rounded with
the nucleus near the center, that of muricatus is
more oval with the nucleus near the side.
The many similarities between rugosus and
grandinatus together with their spatial isolation
suggest the possibility that they may have
evolved from the same species stock. Although a
subspecific relationship may be indicated, the
two are here considered to have developed full
specific differentiation.
Habitat—Usually occurs on low islands within
the geographic range; on coral reef flats near the
high tide line, on jagged pieces of raised reef.
Description—Shell reaching 37.9 mm (about 1
'/l> inches) in length, elongate conical in shape,
average obesity about .72 (44 specimens range
from .62-.82); rather solidly and thickly con-
structed for its size, imperforate; mature speci-
mens with very deeply impressed suture and
rounded whorls; sculptured with four spiral rows
per whorl of stubby spines. External shell color
yellowish white; often coated with what appears
Littorinidae
to be a light-yellow to brown or rather dark
grayish brown periostracum which may wear thin
especially at tips of spines; no definite color
pattern apparent; occasional lines or splotches of
medium brown present; upper (most posterior)
part of aperture usually covered with a brown
glaze. Base moderately flattened, sculptured spi-
rally with nodulose cords. Whorls 7-9, moderately
rounded. Length of spire usually considerably
greater than half the length of shell. Spire convex,
produced at an angle of from 58-60°. Aperture
rounded; outer lip considerably thickened, pli-
cate within, tapering to a thinner, crenulate edge.
Inner lip with a thin, brown glaze posteriorly,
forming a tooth-like bulge anteriorly, near junc-
tion with outer lip near base of columella. Suture
often deeply impressed, typically forming a chan-
nel between whorls. Primary sculptural feature is
the four spiral rows of spines. Spines not regu-
larly aligned axially, although anteriormost two
rows more-so than others; from about 17-25
spines per row on body whorl; bases of anteri-
ormost 3 rows of spines joined by low spiral
carinae; bases of posteriormost row of spines
usually separate; second from anterior row usual-
ly the smallest. Secondary spiral sculpture be-
Plate 393. Tectarius grandinatus (Gmelin, 1791).
Figs. 1,2. Trochus grandinatus Gmelin, lectotype in ZMC,
specimen figured by Chemnitz, Conchylien Cabinet, vol.
10, pl. 169, fig. 1639, from Palmerston Atoll, Cook Is-
lands, 32 x 22.4 mm.
Figs. 3-5. Monodonta coronaria Lamarck, holotype,
MHNG 1096/23, 41 x 27 mm.
[05-452]
January 15, 1972
tween rows of spines consisting of 2-4 raised
cords; overall microscopic sculpture of fine,
closely-spaced spiral threads. Axial sculpture
consisting of fine, irregular, closely-spaced, over-
lapping lines of growth. Shell surface under high
magnification may show minute closely-spaced
perforations. Operculum moderate in size,
rounded-oval, an average one measuring about 9
mm. in diameter, thin, light-brown with a dark-
brown center, paucispiral, nucleus about central.
A thin, light- to dark-brown periostracum present;
closely applied but easily worn away; perios-
tracum smooth and dully shining. Nuclear whorls
about | !/2, smooth and shining; first post-nuclear
whorl rapidly becoming multi-carinate and de-
veloping spines on the second. Radula littorinoid,
2-1-1-1-2; lateral tooth with a vertical partition
and with an embayment characteristic of Lit-
torinidae. Animal moderately large, littorinoid.
Penis fairly large and apparently quite extensible;
seminal groove in deep fold running along medial
edge to tip and bordered by thickened, papillose
glandular-appearing tissue; distal end of penis
vermiform; lateral edge of penis lined with large
number of glands not extending onto vermiform
tip. Reproductive data and life history unknown.
Measurements (mm)—
locality
length width no. whorls
BYR) 23.4 7+ Manihi, Tuamotu Ids.
34.3 D5 T+ Aitutaki, Cook Ids.
32.0 22.8 6+ Mangaia, Cook Ids.
29.8 9519 7+ Mangareva, Gambier Ids.
94.3 18.7 S+ Bird Id., Palmerston Atoll]
22.0 17.0 9 Bird Id., Palmerston Atoll]
20.7 15.7 7+ Cooks Motu, Palmerston Atoll
18.1 14.7 7+ Mangaia, Cook Ids.
16.1 132 6+ Aitutaki, Cook Ids.
14.3 10.8 6+ Aitutaki, Cook Ids.
12.4 9.1 9 Tikahau Atoll, Tuamotu Ids.
7.8 6.0 6+ Mangaia, Cook Ids.
Synonymy—
[1784 Trochus bullatus Martyn, The Universal Conchologist,
vol. 1, fig. 38; rejected work, I.C.Z.N. Opinion 456].
1791 Trochus grandinatus Gmelin, Systema Naturae, ed. 13,
p. 3585 (ad Palmerstoni insulam [=Palmerston Atoll,
Cook Islands); refers to Chemnitz “Conch” vol. 10, p.
291, pl. 169, fig. 1639 and to Martyn “Conch”, vol. 1,
fig. 38. Lectotype, here selected, specimen from Spen-
gler Collection, ZMC, figured by Chemnitz; see our
pl. 393, figs. 1 and 2, 32 x 22.4 mm.
1816 Monodonta coronaria Lamarck, Liste Des Objets Repré-
sentés, Tableau Eneyclopédique et Méthodique, part
23, p. 10, pl. 447, fig. 6 a,b (no locality given);
Holotype MHNG 1096/23, 41 x 27 mm; 1822, His-
toire Naturelle Des Animaux sans Vertébres, vol. 7,
Deooe
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
i
Tectarius 51
Plate 394. Tectarius grandinatus (Gmelin).
Fig. A. Radula.
Fig. B. Penis (both from Palmerston Atoll, Cook Islands,
USNM 685165).
Types—The lectotype of Trochus grandinatus
Ginelin, the specimen figured by Chemnitz, vol.
10, p. 291, pl. 169, fig. 1639, is in the Zoological
Museum Copenhagen. It measures 32 x 22.4 mm
(see pl. 393, figs. 1,2). The Holotype of Monodon-
ta coronaria Lamarck is in the MHNG 1096/23
(see pl. 393, figs. 3-5). It measures 41 x 27 mm.
180° 160° 140°
| T T T XX
\
midway)» Pearl Hermes
Haw,
Alia
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a
Oohu O3,
Woke Pa ‘s ; :
|
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SLHEBRIDES
Mois
k/ Sate FIJNis
Kes
NeW bia
EDONIA. = oe es
. Norfolk
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Plate 395. Geographic distribution of Tectarius grandinatus
(Gmelin) in the southeastern Pacific Ocean.
[05-453]
518 Joseph Rosewater
Littorinidae
Records—COOK ISLANDS: Palmerston Id.; Aitutaki (both
USNM, ANSP, Del. Mus. N.H.); Hervey Ids. (USNM, MCZ,
ANSP); E. side Koromiri Id., S.E. Rarotonga (ANSP); Mangaia
(USNM). SOCIETY ISLANDS: W. coast Bora Bora (ANSP);
Tahiti (USNM, MCZ, ANSP). TUAMOTU ISLANDS: Manihi
Atoll (USNM); Takaroa Atoll (ANSP); Maiai Id., Tikehau
Atoll (USNM); Aratika Atoll (ANSP); N. of Temao Harbor,
Makatea Id. (USNM); Toau Atoll; Raroia Atoll; (all ANSP)
Raraka Atoll (Del. Mus. N.H.); Fakarava Atoll; Makemo Atoll;
Tatatkoto Atoll; Anaa Atoll; Vahitahi Atoll; Nengonengo
Atoll; Tureia Atoll (all USNM); Marutea Atoll, Acteon Group
(ZMA). GAMBIER ISLANDS: Mangareva Id. (USNM, MCZ).
Tectarius pagodus (Linné, 1758)
(Pl. 388, figs. 5-7
Range—From the Philippines, through the
Western Pacific Are to the Solomon Islands.
Remarks—The name applied by Linné to this
species could not have been more appropriately
descriptive as the shell with its usually upturned
spines very much resembles an oriental turreted
pagoda. As pointed out by Dodge (1959) there are
a number of similarities between T. pagodus and
T. tectumpersicum, both described by Linné.
However pagodus reaches a larger size, 21/2
inches versus 1 inch, is proportionately broader
and has more prominent spinose carinae than
tectumpersicum. There is seldom any difficulty in
separating these species with comparative mate-
rial at hand.
The method of reproduction in pagodus re-
mains to be observed. The normal habitat of the
species is on cliffs above the sea. As there is no
evidence that the species is ovoviviparous, there
must be a periodic migration to the sea for breed-
ing purposes, although such a phenomenon has
not been reported.
Another question arising from the high shore
habitat of these nominally marine snails concerns
their food. Rumphius (1705) early suggested that
they feed upon the cliffs where they live. Ex-
amination of some fecal pellets of this species
revealed the presence of considerable debris and
what appeared to be plant cells. It is quite likely
that T. pagodus feeds on plant life such as algae
and lichens growing on the sea cliffs. The radula
is extremely long which may denote such a
browsing manner of feeding, the extra length
possibly being required because aerial feeding
without lubrication from sea water causes a rapid
wearing of the teeth (see Quoy and Gaimard,
Astrolabe, pl. 62, fig. 1).
Habitat—*‘. ... these animals suck their food
from the briny moisture of the cliffs to which they
cling, being unable to endure the water”
(Rumphius, 1705, p. 74). Found on vertical lime-
stone cliffs 1-2 meters above high tide line (per-
sonal observations, Moluccas Islands, Indonesia,
1970).
Description—Shell reaching 61.4 mm. (about
2'/o inches) in length, squatly conical in shape,
average obesity about .93 (18 specimens range
from .84-1.1); mature individuals rather heavily
constructed, imperforate, and sculptured on the
body whorl with 2 carinate rows of usually thick,
straightly projecting or slightly upturned spines,
between which spiral cords are roughly produced
and the shell surface often thrown into oblique
waves. Overall external color yellowish to grayish
white, with no discernable pattern; in young
specimens dark-brown spiral lines may be pres-
ent externally or within aperture; aperture usually
yellowish brown, its edge white. Base distinctly
flattened, sculptured spirally with nodulose cords
extending into aperture; base separated from
upper part of body whorl by one of the rows of
spines at periphery. Whorls 5-8, flat-sided, ex-
cepting spines. Length of spire usually greater
than half the length of shell. Spire convex, pro-
duced at an angle, excepting spines, of from
59-65°. Aperture broadly rounded; outer lip thick-
ly produced in mature individuals, strongly pli-
cate within; plicae not reaching edge of aperture;
inner lip smooth, white posteriorly, but forming a
prominent tooth-like bulge anteriorly near junc-
tion with outer lip at base of columella. Suture
somewhat obscure, marked by protrusion of sec-
ondary carinae of succeeding whorl. Outstanding
sculptural feature is the midwhorl spinose carina,
with typically from 9-12 antero-posteriorly flat-
tened, triangularly-shaped spines on carina of
body whorl. Surface of shell at base of spines
raised to form oblique fold often reaching to
suture. Secondary spine bearing carinae at per-
Plate 396. Tectarius pagodus (Linné, 1758).
Fig. 1. Turbo pagodus Linné; lectotype figure from Argen-
ville, pl. 11, fig. a.
Figs. 2,3. Monodonta bicolor Lamarck, holotype, MHNG
1096/19, 54 x 38.2 mm.
[05-454]
January 15, 1972
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
Tectarius 519
iphery of each whorl, entirely visible only on
body whorl and protruding above suture of pre-
ceeding whorls. Secondary spiral sculpture of
raised, roughened spiral cords in turn, covered
with countless closely-spaced spiral microscopic
textural threads. Operculum large (average about
16 mm. diameter) thin, brown, circular, corneous,
paucispiral, nucleus about central. Periostracum
not evident. Nuclear whorls partially decollate in
all specimens examined, earliest whorl (probably
first postnuclear) is spirally striate, the midwhorl
carina beginning about second postnuclear
whorl; carina becoming nodulose or prespinose
almost immediately. Radula littorinoid (2-1-1-1-2)
extremely long; lateral tooth partitioned and with
an embayment. Animal large, also littorinoid;
sides of foot and tentacles bright yellowish
orange, remainder of animal grayish brown (color
observations on living animals from Kai Islands,
Moluccas, Indonesia); penis well-developed,
with the seminal duct contained within a deep
fold running along its medial edge; with a large
number of glands attached along lateral edge;
penis minutely papillose also over its surface,
unbranched. Reproductive data and life history
unknown.
Measurements (mm) (width includes spines)
locality
length width — no. whorls
61.4 45.0 7+ Lutee, Choiseul Id.
Solomon Islands
57.1 47.8 6+ Pavuvu Id., Russel
Group, Solomon Islands
49.3 42.3 7+ Lutee, Choiseul Id.
Solomon Islands
46.9 43.4 8+ Pavuvu Id., Russel
Group, Solomon Islands
44.4 42.3 7+ Bougainville Id.,
Solomon Islands
41.3 37.4 5+ Majugag Id., W coast
Buka Id., Solomon
Islands
39.7 40.0 8 Lutee, Choiseul Id.,
Solomon Islands
31.5 31.9 6+ Balagnan Id., Surigao
District, Mindanao,
Philippines
275 25.5 8 Soepiori Ids., Schouten
Ids., West Irian
14.3 12.6 Hf Biak, West Irian
Synonymy—
1758 Turbo pagodus Linné, Systema Naturae, ed. 10, p. 762;
refers to Argenville, pl. bl, fig. A “Pagodus”’ [lecto-
type figure]; (type-locality, O. Asiatico, here restricted
to Amboina, Moluccas).
1822 Monodonta bicolor Lamarck, Animaux san Vertébres,
vol. 7, p. 31 (no locality); holotype in MHNG 1096/19,
54 x 38.2 mm.
1840 Pagodella major Swainson, A Treatise on Malacology,
p. 351 (no locality given); refers to Chemnitz, pl. 163,
figs. 1541, 1542 (specimen figured is lectotype, pos-
sibly in Copenhagen Museum).
1850 Pagodus verus J. E. Gray in M. E. Gray, Figures of
Molluscous Animals, vol. 4, p. 78; refers to Trochus
pagodus Quoy [and Gaimard, Astrolabe] pl. [62, not]
“82”. figs. 1-4; not Littorina papillosa var. vera Phil-
ippi, 1846 [= Tectarius tectumpersicum Linné].
Types—The location of Linné’s type of Turbo
pagodus is unknown, and Dance (1967) has
pointed out that this species is missing from the
Linnaean collection in London. In the absence of
a type, one of the figures cited by Linné in
connection with the original description is here
Plate 397. Tectarius pagodus (Linné).
Fig. A. Radula of specimen from West Irian; note smooth
cusps probably denoting worn teeth, also “partitioned”
lateral.
Fig. B. Penis, anterior, and C. posterior views respectively;
note glands on lateral edge and papillose surface (both
ANSP 207638).
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Plate 398. Geographic distribution of Tectarius pagodus
(Linné) in the central Indo-Pacific faunal region.
[05-455]
520 Joseph Rosewater
designated as representing the lectotype: Argen-
ville, pl. 11, fig. A. The type-locality, originally
given as “QO. Asiatico” is here restricted to Amboi-
na, Moluccas. The holotype of Monodonta bicolor
Lamarck is in the Museum d’Histoire Naturelle,
Geneva (MHNG register no. 1096/19). The loca-
tion of types of Swainsons’ Pagodella major and
of Gray’s Pagodus verus are unknown to me, and
the figures cited in connection with their descrip-
tions may be considered as representing their
lectotypes (see Synonymy).
Records—PHILIPPINES: E. coast Polillo (Del. Mus. N.H.,
USNM): Calapan, Mindoro Id. (Del. Mus. N.H.); Balagnan
Id., Surigao District (USNM); Zamboanga, both Mindanao
(ANSP Del. Mus. N.H.). INDONESIA: Pulau We, Sumatra
(RNHL); Java (ANSP, RNHL); Timor (RNHL); Morotai Id.;
Toetoe Id.; Dagaseli, both N Loloda Group (all MCZ); Ter-
nate (RNHL); Buru Id. (ZMA); Ambon (MCZ, ZMA): Tanim-
bar Islands (RNHL); Kur Id.; Warbal Id., W of Nuhu Rowa,
both Kai Ids. (both USNM, WAM). NEW GUINEA: Waigeo
Id. (ANSP); Misool Id.; Fakfak (both Leiden); Manokwari
(ANSP); Biak; Soepiori Id., both Schouten Ids. (both USNM);
Rouw, Aoeri Ids. (ANSP); Woodlark Id. (MCZ). SOLOMON
ISLANDS: Majugag Id., W coast Buka Id.; Nr. Kihili, Buin,
Bougainville Id. (both USNM); Choiseul Bay (ANSP); Lutee,
both Choiseul Id. (ANSP, USNM): Ataa District, Malaita
(ANSP); Roviana (MCZ); Pavuvu, Russell Group (USNM).
Tectarius tectumpersicum (Linné, 1758)
(Pl. 388, figs. 3,4)
Range—From the Philippines along the West-
ern Pacific Arc through Melanesia.
Remarks—Tectarius tectumpersicum is grossly
similar in many ways to T. pagodus and it often
appears difficult to construct a point by point
comparative description which clearly differen-
tiates the two species except in matters of size and
degree of obesity; pagodus reaches a length of
21/2 inches (61 mm) while tectumpersicum rarely
reaches 1'/2 inches (34 mm); pagodus is very
obese, sometimes being wider than high, but in
tectumpersicum the width of shell is generally
only about 77% of the length. Differences are
apparent also in spinosity, there being two rows
of spines on the body whorl of pagodus and 3
rows in tectumpersicum. Spines in tectumpersi-
cum tend to be stubby and round-ended while in
pagodus they are pointed and broadly triangular,
although there are occasional specimens of tec-
tumpersicum which tend to resemble small, ma-
ture pagodus. Generally, however, specimens of
pagodus the size of tectumpersicum are obvious-
ly immature and thin-lipped, so that the “rule-of-
thumb” involving size of specimens can be de-
pended on to separate the species.
Littorinidae
Habitat—Lives in pockets of worn, raised lime-
stone reef rock, 1-2 meters above high tide line
(personal observations, Moluccas Islands, In-
donesia, 1970).
Description—Shell reaching 34.6 mm (about
1*/16 inches) in length, conical in shape, average
obesity about .77 (38 specimens range from .64
~.85), mature individuals moderately thick in
structure, imperforate, and sculptured with three
main rows of stubby, often upturned, spines on
body whorl, and usually with two rows on spire
whorls, between which spiral cords are roughly
produced, wavy or papillose and often approach-
ing minor rows of spines. External color yellow-
ish to grayish white, with no regular patterning
although some specimens have diffuse dark spiral
color bands externally or within aperture es-
pecially at edge of outer lip or on tooth-like bulge
of inner lip. Remainder of aperture usually white
or yellowish white. Base moderately flattened,
Plate. 399. Tectarius tectumpersicum (Linné, 1758).
Figs. 1-3. Turbo tectumpersicum Linné, lectotype in Lin-
nean Society of London collection, ca. 25 x 23 mm.
Figs. 4-7. Monodonta papillosa Lamarck, lectotype,
MHNG 1096/22-2, 29.3 x 25.5 mm.
[05-456]
January 15, 1972
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
Tectarius 52]
Plate 400. Tectarius tectumpersicum (Linne).
Fig. A. Radula.
Fig. B. Penis (both from Biak, West Irian, ANSP 206421
and USNM 637390).
sculptured spirally with nodulose cords which
may extend into aperture; base separated from
upper part of body whorl by a row of spines at
periphery. Whorls 6-8, rather flatsided excepting
spines. Length of spire greater than half the
length of shell. Spire convex, produced at an
angle of from about 55-65°. Aperture compactly
rounded; outer lip thickly produced; strongly
plicate within, but thin and crenulate at apertural
edge; inner lip smooth posteriorly, but forming a
tooth-like bulge anteriorly near junction with
outer lip at base of columella. Suture obscure,
partially masked by lower row of spines at pe-
riphery of whorls. Center row usually bearing
largest spines, from 9-12 on body whorl; the more
posterior row, near suture has smaller spines but
about same number; anterior row at periphery
smaller still and more numerous with from 13-16
spines. On spire whorls center row of spines may
predominate with others being either hidden or
suppressed. Spines usually not aligned axially.
Secondary spiral cords in some specimens almost
as spinose as primary ones; in other specimens
hardly noticeable. Entire surface covered with
closely spaced spiral microscopic — textural
threads. Axial sculpture consists of often coarse,
irregular lines of growth. Operculum moderate in
size (average about 6-7 mm diameter), thin,
brown, circular, corneous, paucispiral, nucleus
about central. Periostracum not evident. Nuclear
whorls about two, but at least partially decollate
in all specimens examined; brown, smooth for at
least 1 volution, then becoming carinate; first post
nuclear whorl weakly nodulose and rapidly be-
coming spinose. Radula littorinoid (2-1-1-1-2)
[teeth about !/2 the size of those of T. pagodus];
lateral tooth partitioned and with an embayment.
Animal medium-sized, littorinoid; penis large,
unbranched, with a large number of glands along
lateral edge, 3/4 the length to tip; surface of penis
otherwise papillose; seminal duct deeply folded.
Life history unknown.
Measurements (mm) (width includes spines)—
locality
length width no. whorls
34.6 24.5 6+ Pavuvu Id., Russell Group,
Solomon Islands
31.4 Bo 8 Lunga, Guadalcanal,
Solomon Islands
96.7 14 7+ Timor, Indonesia
22.8 18.4 6+ Cebu, Philippines
21.4 15.9 7+ Biak, West Irian
20.0 5:5 6+ Biak, West Irian
19.2 16.0 6+ Cebu, Philippines
18.1 13:7 7+ Biak, West Irian
15.2 12.9 6+ Anir Id., New Ireland
13.6 10.3 8 Philippines
Synonymy—
1758 Turbo tectumpersicum Linné, Systema Naturae, ed. 10,
p. 762 (no locality given; Cebu Id., Philippines, here
selected as type-locality); lectotype in Linnean Soci-
ety of London collection.
—— Trochus bullatus “Martvn” of authors; [not T. bullatus
Martyn, 1784, Universal Conchologist, vol. 1, fig. 38;
non-binomial; is Tectarius grandinatus (Gmelin,
1791)].
1822 Monodonta papillosa Lamarck, Histoire Naturelle des
Animaux sans Vertébres, vol. 7, p. 32. (“les mers de
Timor’); lectotype MHNG 1096/22-2.
1846 Litorina papillosa vera Philippi, Abbildungen und Be-
schreibungen Conchylien, vol. 2, p. 141 (no locality
given); refers to “Delessert, Recueil, pl. 36, fig. 10,”
here selected as the lectotype figure.
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Plate 401. Geographic distribution of Tectarius tectumper-
sicum (Linne) in the central Indo-Pacific faunal region.
[05-457]
O22 Joseph Rosewater Littorinidae
Types—The specimen of Turbo tectumper- in structure, imperforate, and sculptured with 2-3
sicum Linné in the Linnaean shell collection
located in the Linnean Society, London, may be
considered the lectotype of this species; it mea-
sures approximately 25mm length (about | inch).
The lectotype somewhat resembles T. pagodus
but may be recognized by its rounded spines and
the mature, thickened shell. The figure cited by
Linné, “Argenville, pl. 11, fig. P’’, is controversial
and a poor representation of the species (also see
Dodge, 1959, p. 229). As no type-locality was
given by Linné, Cebu Island, Philippines, is here
selected as a place from which specimens could
have come in the early 18th century. The figure
referred to by Philippi, “Delessert, Recueil, pl.
36, fig. 10”, is here selected as the lectotype figure
of Litorina papillosa vera. The lectotype of Mon-
odonta papillosa Lamarck, which proves to be an
absolute synonym of T. tectumpersicum is in the
Geneva Museum (MHNG 1096/22-2; see our pl.
399 figs. 4-7).
Nomenclature—The tectumpersicum
does not need to agree in gender with Tectarius
because the former is a noun in apposition, mean-
name
ing “persian roof”.
Records—PHILIPPINES: Cebu (USNM). INDONESIA:
Timor (USNM, RNHL); Ambon; Kur Id.; Warbal Id., W of
Nuhu Rowa, both Kai Islands (all USNM, WAM). NEW
GUINEA: Wasior, Wandammen Bay (ZMA); Soepiori Id.,
(MCZ): Biak Id., both Schouten Islands, all West Irian
(USNM); reef at Cape Moem, nr. Wewak, Territory of New
Guinea (MCZ). BISMARCK ARCHIPELAGO: Feni Ids., E of
New Ireland (USNM). SOLOMON ISLANDS: Nissan Id.,
Green Islands (SMF); Choiseul Bay, Choiseul Id. (ANSP);
Stirling Isle, Treasury Ids.; Munda, New Georgia; Pavuvu Id.,
Russell Group; Lunga, Guadalcanal (all USNM); Santa Ana
Id., S. of San Cristobal (ANSP).
Tectarius rusticus (Philippi, 1846)
(Pl. 388, figs. 8,9; pls. 402, 403)
Range—Northern Western Australia.
Remarks—This species is restricted to northern
Western Australia according to available locality
data. It appears to be most closely related to T.
rugosus trom which it differs in its usually less
regularly sculptured, more flat-sided whorls and
in its lack of external coloration. In T. rusticus the
subsutural, midwhorl and peripheral rows of
spines are quite commonly strongly expressed,
while in rugosus all rows are subequal.
Habitat—Lives on rocks above high tide line.
Description—Shell reaching about 40 mm
(about 1°/s inches) in length, broadly conical in
shape, average obesity about .77 (29 specimens
range from .72-.83); becoming moderately thick
main rows per whorl of rather muted spines.
External color yellowish white with occasional
faint orange stripes; inside of aperture white and
occasionally tinged with pinkish orange, often
with brown lines at its edge and revolving within;
apertural tooth often stained with brown. Base
flattened, sculptured spirally with nodulose
cords, the strongest of which occurs just below
periphery of body whorl. Whorls 6-8, rather flat
sided. Length of spire greater than half the length
Plate 402. Tectarius rusticus (Philippi, 1846).
Figs. 1,2. Litorina papillosa rustica Philippi, knobby and
smooth forms, respectively.
Fig. 2 (lectotype figure) probably is representation of Phi-
lippi’s concept of L. p. subinermis, the smooth form; both
from Point Swan, northern Western Australia; from Ab-
bildungen und Beschreibungen Conchylien, vol. 2,
Litorina, pl. 2, figs. 3,4.
[05-458]
January 15, 1972
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
Tectarius 523
of shell or the two may be about subequal. Spire
convex, produced at an angle of about 68°. Aper-
ture rounded to squarish; outer lip thickly pro-
duced, internal plicae only moderately produced
and not reaching edge of aperture; outer lip
tapering to thin crenulate edge; inner lip forming
a tooth-like bulge anteriorly near junction with
outer lip; tooth often stained with brown. Suture
usually obscured by anteriormost row of spines of
preceding whorl. Spiral spinose sculpture sub-
dued, usually three rows are outstanding: the
anteriormost, posteriormost and central rows of
each whorl, the rest being limited to undulating,
slightly bumpy cords; from 12-14 spines per row
on body whorl; spines sometimes coalesce into
oblique axial bars. Fine sculpture composed of
microscopic wavy spiral cords and finer threads.
Operculum moderate in size, circular, light-
brown with a dark-brown center, paucispiral,
nucleus about central. Periostracum not evident.
Nuclear whorls partly decollate in all specimens
examined; remaining portions smooth, white;
early postnuclear whorls rapidly becoming spi-
rally striate. Radula littorinoid, 2-1-1-1-2, similar
in appearance to that of T. rugosus. Preserved
specimens not available for observations on anat-
omy; radula obtained from dried specimen. No-
thing reported concerning reproduction and de-
velopment.
Measurements (mm) (all Western Australia)—
length width no. whorls locality
35.5 29.0 5+ Yampi Sound, W.A.
34.5 D7A5 6+ Cliff Id., King Sound
32.4 2319 8+ Buccaneer Archipelago
25.8 20.6 7+ Buccaneer Archipelago
D322; 18.8 5+ Cliff Id., King Sound
22.1 17.1 6+ Buccaneer Archipelago
21.9 16.2 7+ Buccaneer Archipelago
21.1 15.3 7+ Cliff Id., King Sound
16.3 12h 6+ Buccaneer Archipelago
15.8 1:22, 6+ Buccaneer Archipelago
Synonymy—
1846 Litorina papillosa rustica Philippi, Byeuiluaen und
Beschreibungen Conchylien, vol. 2, Litorina, p. 140,
pl. 2, fig. 3 [and 4 Lectotype Fe (Point Swan
[Cape Leveque, Dampier Land, northern Western
Australia]; type-specimen may be in Berlin Museum).
1846 Litorina papillosa subinermis Philippi, ibid., p. 141. pl.
2, fig. 4
1857 Littorina bullata in Reeve, Conchologia Iconica, vol. 10,
Littorina, pl. 1, fig. lc; not Trochus bullatus Martyn
[non-binomial] which is Tectarius grandinatus
Gmelin.
1971 Tectarius pagodus Linnaeus, Wilson and Gillett, Aus-
tralian Shells, p. 30, pl. 11, fig. 9.
Records—WESTERN AUSTRALIA: Troughton Islands, N.
a Admiralty Gulf; S.E. Wood Island, E of Cockatoo Island:
Yampi Sound; Koolan Island (all WAM); Kellan Island, Buc-
caneer Archipelago (AMS; USNM; MCZ): Cliff Island
(USNM) all vicinity of King Sound.
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Plate 403. Geographic distribution of Tectarius rusticus
(Philippi) in northern Western Australia.
Tectarius songoense (Martin, 1931)
(Pl. 404, figs. 3,4)
Range—Eocene of Java, Nanggoelan-beds, Kali
Songo.
Remarks—It is not too difficult to determine
from Martin’s illustration just why he assigned
this species to “Tectarium”. There is a weakly
nodulose sculpture, perhaps better described as
beaded, and there appears to be a columellar
tooth, a structure characteristic of Tectariinae.
The general shape and appearance are, however,
more suggestive of Trochidae. The species is
tentatively here placed in the subfamily Tectari-
inae for want of positive proof to the contrary.
Synonymy—
1931 Tectarium (Echinella) songoense kK. Martin, Wetens-
schappelijke Mededeelingen Dienst Mijnbouw, no.
18, p. 41, pl. 6, fig. 5, 5a, (Upper Eocene, Nanggoe-
lanbeds, Java); unique holotype probably in Geologi-
cal Museum, Leiden; length $8 mm.
[05-459]
524 Joseph Rosewater
Littorinidae
Subgenus Subditotectarius Ladd, 1966
Type: Tectarius rehderi Ladd, 1966
Small, stout; spire conical, base convex; with
beaded spiral ribs and slightly oblique axial lines;
aperture strongly plicate within. Sculpture sub-
dued, consisting of small beads formed by con-
vergence of axial growth lines and spiral threads.
Synonymy—
1966 Subditotectarius Ladd, Geological Survey Professional
Paper 531, p. 59; type-species by original designation:
Tectarius rehderi Ladd.
Tectarius (Subditotectarius) rehderi Ladd, 1966
(Pl. 404, figs. 5-7)
Range—Lower Miocene, Marshall Islands.
Remarks—The fossil Subditotectarius rehderi
is characterized by its subdued beaded sculpture
as compared with Tectarius s.s. with its larger
nodules and spines. Otherwise the species is
quite similar to other Tectarius in outline and
sculpture, and in the presence of a columellar
tooth and plications within the aperture. Al-
though apparently clearly a tectariine S. rehderiis
not believed to be closely related to any living
species.
Synonymy—
1966 Tectarius (Subditotectarius) rehderi Ladd, Geological
Survey Professional Paper 531, p. 59, pl. 11, figs.
11-13 (drill hole 2A, Bikini Atoll, 1,051-1,057 feet;
early miocene); holotvpe USNM 648342, 2.8x 2.4mm.
Plate 404. Fossils of Tectariinae and Echinininae.
Figs. 1,2. Echininus adelaidensis (Cotton, 1947) holotype,
16 x 12 mm.
Figs. 3,4. Tectarius songoense Martin, 1931; holotype, 8
mm length.
Figs. 5-7. Tectarius rehderi Ladd, 1966, holotype, 2.8 « 2.4
mim.
[05-460]
January 15, 1972
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
Subfamily Echinininae, new subfamily
Genus Echininus Clench and Abbott, 1942
Type: Echininus cumingi (Philippi, 1846)
The genus Echininus sensu lato includes the
nominate subgenus, Echininus, typified by the
umbilicate and spinose EF. cumingi and also E.
cumingi spinulosus, both of the western Pacific,
and the subgenus Tectininus Clench and Abbott,
1942, having the non-umbilicate and less spinose
monotype, E. (T.) nodulosus (Pfeiffer), a species
which inhabits the Caribbean area. Members of
both subgenera have what may be called multi-
spiral (polygyrous spiral type) opercula and pos-
sess moderately spinose shells. In Echininus
sensu stricto the central radula tooth is moder-
ately reduced in width, while in Tectininus it is
dramatically reduced and narrowed. Abbott
(1954) discussed the phylogenetic position of
Echininus and concluded that it belongs in Lit-
torinidae although it possesses many specialized
characters such as the multispiral operculum, an
umbilicus and a narrow central radula tooth. As
there are no representatives of Tectininus in the
Indo-Pacific, only Echininus s.s. will be con-
sidered here.
Synonymy—
1854 Echinella ‘Swainson’ H. and A. Adams, The Genera of
Recent Mollusca, vol. 1, p. 316; three species men-
tioned: coronaria Lamarck [= Tectarius grandinatus
Gmelin]; granulata Swainson [unrecognizable]; and
the exemplary species, cumingii Philippi; 1895, Pils-
bry, Catalogue of the Marine Mollusks of Japan, p.
175; 1901, Pilsbry, Proceedings of the Academy of
Natural Sciences of Philadelphia, p. 198; 1903, Kes-
teven, Proceedings of the Linnean Society of New
South Wales, for 1902, part 4, p. 632, in part; not
Echinella Swainson, 1840 [= Tectarius s.s.].
1942 Echininus Clench and Abbott, Johnsonia, vol. 1, no. 4,
p. 3; new name for Nina Gray, 1850, Figures of
Molluscous Animals, London, vol. 4, p. 78; Type
species by monotypy Trochus cumingii Philippi; not
Nina Horsfield, 1829, nor Gray, 1855.
Subgenus Echininus s.s.
Pyramidal, umbilicate littorinids with a stron-
gly spinose shell. In males the penis has a deep
but open sperm duct, and basal penial glands are
present. Operculum multispiral. Radula littori-
noid, the central tooth moderately narrow.
Echininus 525
Echininus cumingi cumingi (Philippi, 1846)
(Pl. 388, figs. 12,13)
Range—From the Philippines along the west-
ern Pacific arc to New Hebrides and the Cook
Islands.
Remarks—At first examination one finds it sur-
prising that “Cuming’s Echininus” is included in
the Littorinidae. This species has many of the
attributes of certain other families, perhaps re-
sembling most some members of the family Tro-
chidae with its conical shape, multispiral opercu-
lum and well-defined umbilicus. Animal charac-
ters, especially the radula, and gross reproductive
features leave no doubt, however, that E. cumingi
is a littorinid. Its closest relative is the smaller
and less spinose, but otherwise very similar ap-
pearing subspecies, EF. cumingi spinulosus (Phil-
ippi). The next most closely related and only
other species in the subfamily, the Western Atlan-
tic E. (Tectininus) nodulosus (Pfeiffer) differs in
never being umbilicate and in having a consider-
ably more reduced radula. The open condition of
the spines occurs occasionally in E. nodulosus
and their arrangement or alignment is similar.
Both have multispiral opercula. Penial anatomy
differs, however, E. cumingi having a cluster of
basal penial glands, whereas E. nodulosus dis-
plays a basal swelling and a single gland located
one half to two thirds the distance to the tip (see
Abbott, 1954, fig. 55, p,q).
Habitat—Lives in pockets of weathered, raised
limestone reef, 3-7 meters above high tide line
(personal observations in Davao, Philippines and
Moluccas Islands, 1970).
Description—Shell reaching 20.4 mm (about 3/4
inch) in length, broadly conical in shape, with
Plate 405. Echininus cumingi cumingi (Philippi, 1846).
Trochus cumingii Philippi, lectotype figure of specimen in
BM(NH), from Reeve, 1857, Conchologia Iconica, vol. 10,
Littorina, pl. 2, fig. 8.
[05-467]
526 Joseph Rosewater
Littorinidae
projecting spines, average obesity about .88 (33
specimens range from .77 to 1.03) only moder-
ately thick in structure, umbilicate; suture im-
pressed, whorls slightly rounded; sculptured
with three rows of short, projecting, often un-
closed spines. External shell color grayish tan, the
spines often being reddish brown; a thin grayish
brown periostrical coating apparent; aperture a
diffuse yellowish to reddish brown, occasionally
with three reddish brown color bands reflecting
the position of the external spines. Base distinctly
flattened, spirally sculptured with nodulose cords
of which about the third below the periphery is
the strongest. Umbilicus very deep but narrow in
mature specimens; bordered medially by a rim of
the columellar callus and distally by a basal fold.
Whorls 6-8, only slightly rounded. Length of
spire greater than half the length of the shell.
Spire convex, produced at an angle of from about
66-76°. Aperture rounded, outer lip only moder-
ately thickened, smooth within, the edge often
undulating in the vicinity of the rows of spines;
inner lip curved and shining, edentulous. Suture
usually somewhat obscured by anteriormost row
of spines of preceding whorl. Primary sculptural
feature is the three spiral rows of spines. Posteri-
ormost row of spines consisting of low, rounded
protuberances; anteriormost row small, unclosed,
moderately projecting, narrow-hoodlike and
closely spaced; middle row of spines largest, most
projecting, also incompletely closed. Rows. of
spines not aligned axially; spine count as follows:
anterior row, 18-25; middle row, 14-17; posterior
15-16 (approximate range). Secondary spiral
sculpture between rows of spines consists of from
4-6 slightly raised cords; overall microscopical
spiral sculpture of fine, closely spaced threads.
Axial sculpture consists of fine irregular often
overlapping lines of growth. Operculum moder-
ate in size, multispiral (polygyrous spiral type)
having about 5-7 volutions, chitinous, dark-
brown (pl. 389). Nuclear whorls smooth, light
brown, shining, about 1.5 volutions; postnuclear
whorls rapidly becoming striately sculptured and
then spinose. Radula littorinoid, 2-1-1-1-2, central
tooth very narrow, rather simple; lateral tooth
only moderately narrow and with a well devel-
oped littorinoid notch.
Animal littorinoid; penis large and_ well-
developed, with a bulbous swelling at its base;
distal extremity simple; with as many as 12 penial
glands clumped mostly on posterior surface near
junction of bulbous base and extremity; sperm
Plate 406. Echininus cumingi (Philippi).
Fig. A. Radula; note “spurs” on central tooth, and generally
narrow dentition.
Fig. B. Penis, 1. anterior, 2. posterior, 3. cross-section of
seminal groove; note cluster of glands near bulbous base
(both from Davao Gulf, Mindanao, Philippines).
groove deeply folded, with an apparent internal
fold best seen distally. Details of reproduction
and life history unknown.
Measurements (mm)—
length width no. whorls locality
20.4 Lio 7 Cook Islands
19.5 17.6 8 Philippines
18.5 14.9 7+ Stirling Isle, Treasury
Ids., Solomons
17.5 14.3 7+ Stirling Isle, Treasury
Ids., Solomons
17.0 14.9 Cook Islands
16.7 15.3 7+ Cook Islands
16.3 14.8 7+ Stirling Isle, Treasury Ids.
Solomon Ids.
15.9 12.3 7+ Stirling Isle, Treasury Ids.
Solomon Ids.
15.1 13.0 7+ Tana, New Hebrides
14.6 13.7 6+ Stirling Isle, Treasury Ids.
Solomon Ids.
13.2 11.9 7+ Stirling Isle, Treasury Ids.
Solomon Ids.
10.6 10.9 8 Philippines
Synonymy—
1846 Trochus cumingii Philippi, Proceedings of the Zoologi-
cal Society of London, for 1845, p. 138 (Guimaras Id.
{south of Panay Id.] Philippines; as Litorina, 1847,
Abbildungen und Beschreibungen Conchylien, vol. 3,
[05-468]
January 15, 1972
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
Echininus 527
Litorina, p. 53, pl. 6, fig. 22; lectotype in BM (NH),
figured by Reeve, 1857, Conchologia Iconica, vol. 10
Littorina, pl. 2, fig. 8.
1879 Trochus echinulatus ‘Kiener’ in P. Fischer, Spécies
Général et Iconographie des Coquilles Vivantes, pl.
43, figs. 2 [name and figure only]; ibid., p. 459, places
name in synonymy of Tectarius cumingi (sic); not
Trochus echinulatus A. Alth, 1850.
Records—PHILIPPINES: Samal Id., Davao Bay, Mindanao
(USNM, MCZ). INDONESIA: Karakelong Id., Talaud Ids.
(MCZ); N. shore Warbal Id., W of Nuhu Rowa, Kai Ids; W.
side Mitak Id., Jamdena Strait, Tanimbar, both Moluccas
(USNM, WAM). NEW GUINEA: Misool; Fakfak (both
RNHL,); Louisade Ids. (RNHL, NMW, ANSP). SOLOMONS:
Stirling Isle, Treasury Ids., (USNM); Santa Ana (ANSP). NEW
HEBRIDES: S end Black Beach, Tana (USNM). COOK
ISLANDS: Mauke, Hervey Ids.; Rarotonga (both ANSP);
Mangaia (USNM).
TUAMor,
Plate 407. Geographic distribution of Echininus cumingi
(Philippi) in the East Indies and Pacific Ocean, and of its more
northerly distributed subspecies E. cumingi spinulosus
(Philippi).
Echininus cumingi spinulosus (Philippi, 1847)
(PI. 388, figs. 14,15; pls. 407, 408)
Range—From southern Japan through the Ryu-
kyu Islands, the northern and western Philip-
pines and eastward to the Mariana Islands.
Remarks—Echininus spinulosus is very close
in its relationship to E. cumingi, the differences
between the two being more of degree than of
kind. The two are apparently geographically iso-
lated, or at least they occupy separate ranges and
so the phenotypic differences may be ecologically
influenced as well as having a genetic basis.
Whatever the basis for the differences between
them, it seems appropriate to consider them as
subspecies. Echininus spinulosus never reaches
as large a size or achieves the squatly conical
shape of E. cumingi, and although the sculpture
of the two is basically very similar, cumingi is
always more distinctly spinose. Both are usually
umbilicate although spinulosus is often narrowly
so and young specimens may lack this feature
entirely, as did Philippi’s type-specimen.
It is interesting to note that E. luchuana was
described by Pilsbry (1901) as a subspecies of
cumingi, which has caused some confusion.
Some malacologists have continued to use the
combination E. cumingi luchuana or simply E.
cumingi when referring to the entity E. spinulo-
sus (Kira, 1959, 1962; Habe, 1951). Philippi’s
figures (1847) clearly show the species concepts
he intended: E. spinulosus (fig. 24), the smaller
less pronouncedly spinose species (which in-
cludes luchuana as a synonym) and EF. cumingi
(fig. 22), the larger and more outstandingly spi-
nose and more squatly conical species.
Habitat—On rocks above the high tide line.
Description—Shell reaching 16.8 mm (about
5/s) inch in length, turbinate in shape, with short
spines; average obesity about .84 (32 specimens
range from .74-1.08) moderately thick in struc-
ture, mature specimens usually umbilicate; su-
ture evident although often obscure; whorls mod-
erately rounded; sculptured with three major
rows of short spines per whorl. External shell
color grayish to tannish orange, the short spines
usually appearing whitish; apertural coloration
Plate 408. Echininus cumingi spinulosus (Philippi, 1847)
Figs. 1-3. Echinella cumingi luchuana Pilsbry, lectotype,
ANSP 70962, 16 x 14 mm.
Fig. 4. Litorina spinulosa Philippi, lectotype figure, from
Abbildungen und Beschreibungen Conchylien, vol. 3,
Litorina, pl. 6, fig. 24.
[05-469]
528 Joseph Rosewater
Littorinidae
tannish to brownish orange, occasionally with 3-4
darker brown color bands revolving within. Base
moderately flattened, sculptured with nodulose
cords of which about the third below the per-
iphery is the strongest. Umbilicus usually rather
narrow and occasionally absent, bordered medi-
ally by a rim of the columella callus and distally
by a poorly to well defined basal fold. Whorls 6-8,
only moderately well rounded. Length of spire
greater than half the length of the shell. Spire
convex, produced at an angle of from about
63-68°. Aperture rounded-oval, outer lip thin to
only moderately thickened, smooth within, and
often slightly undulating in vicinity of external
spine rows; inner lip curved and shining, edentu-
lous. Suture fairly evident to somewhat obscured
by anteriormost row of spines of preceding whorl.
Primary sculptural feature is three spiral rows of
spines located centrally, anteriorly and posteri-
orly on each whorl; spiral cords between main
rows tending to become nodular to spinose. Rows
of spines not well aligned axially; spine count as
follows: anterior row about 24; middle row about
19-23; posterior row about 16-19. Secondary
sculpture consisting of 4-5 rows of spiral cords
which occasionally become nearly as strongly
spinose as the 3 primary rows which they sepa-
rate; overall microscopical spiral sculpture of
fine, closely-spaced threads. Axial sculpture con-
sists of fine irregular lines of growth. Operculum
small to moderate in size, multispiral (polygyrous
spiral type) having 5-6 volutions, chitinous, dark
brown. Nuclear whorls smooth, tannish white,
shining, about 1.5 volutions; postnuclear whorls
rapidly becoming striated and nodulose. Radula
littorinoid, 2-1-1-1-2; central tooth narrow and
reduced; lateral tooth with a distinct littorinoid
notch. Animal littorinoid; penis unbranched; a
well developed sperm groove running along its
medial edge; distal portion papillose; 2-3 penial
glands located half-way between base and tip in
preserved specimen. Nothing has been reported
concerning reproduction and life history.
Measurements (mm)—
length width no. whorls locality
16.8 11.7 8 Fuga Id., Philippines
15.6 12.2 7+ “Japan”
14.9 Val T+ Yokohama, Japan
14.2 11.8 7 Batan Id., Philippines
13.9 11.5 6+ Batan Id., Philippines
13.5 10.9 8 Batan Id., Philippines
13.0 10.4 7 Kume-shima, Ryukyu Ids.
12.6 10.4 6+ Okinawa, Ryukyu Ids.
12.0 10.0 7+ Batan Id., Philippines
11.1 9.0 a Batan Id., Philippines
10.8 10.5 7 Batan Id., Philippines
7.3 7.9 6 Batan Id. Philippines
Synonymy—
1847 Litorina spinulosa Philippi, Abbildungen und Beschrei-
bungen Conchylien, vol. 3, Litorina, p. 53, pl. 6, fig.
24, Lectotype Figure (Manila); type may be in Berlin
Museum.
1895 ‘Echinella cumingi Phil. in Pilsbry, Catalogue of the
Marine Mollusks of Japan, published by Frederick
Stearns, Detroit, p. 175 (Yaeyama [Okinawa)]).
1901 Echinella cumingi luchuana Pilsbry, Proceedings of the
Academy of Natural Sciences of Philadelphia, p. 198
(Loo Choo Islands [Ryukyu Islands]); ibid. p. 394, pl.
19, fig. 16; lectotvpe ANSP 70962, ca. 16 « 14 mm.
Records—JAPAN: Yokohama (USNM). RYUKYU_ IS-
LANDS: Kadena Circle (USNM):; Tsukin-shima, both Okina-
wa (MCZ): Kume-shima (MCZ ANSP, USNM, BPBM); Kari-
mata, Miyako-shima (MCZ, ANSP); Ora Wan (USNM). TAI-
wan; Hung-t?ou Hsu, off Pacific Coast (ANSP, USNM).
PHILIPPINES: Santa Domingo de Basco, Batan, Batan Ids.
(USNM); Dalupiri Id. (MCZ); Fuga Id., both Babuyan Ids.
(USNM); Camp Wallace, Province of La Union, Luzon,
Puerto Princessa, Palawan (both USNM). MARIANA IS-
LANDS: Saipan; Piti Bay, Guam (both ANSP); Apra Bay;
Asan Point, both Guam (both USNM).
Echininus adelaidensis (Cotton, 1947)
(Pl. 404, figs. 1,2)
Range—Adelaidean (Pliocene), South Aus-
tralia.
Remarks— Echininus adelaidensis certainly is
a unique appearing species which somewhat re-
sembles certain of the Trochidae nearly as much
as it does Echininus (cf. Turcica A. Adams or
Perrinia H. and A. Adams, as shown in Wenz,
1938). The characteristics of its partly open
spines and the presence of an umbilicus may be
sufficient to relate it to Echininus, however. Cot-
ton refers to E. cumingi Philippi having been
collected in Caloundra, Queensland and Western
Australia, but I have not seen such records in the
course of the present study. It is possible that
northern portions of Australia may be within the
range of E. cumingi which reaches the southern
Moluccas and New Guinea.
Synonymy—
1947 Nina adelaidensis Cotton, Records of the South Austra-
lian Museum, vol. 8, no. 4, p. 666, pl. 21, figs. 17, 18
(Adelaidean Pliocene, Salisbury Bore, 350 feet); holo-
type in Tate Museum, University of Adelaide, 16 « 12
mm.
[05-470]
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
January 15, 1972
[replacing vol. 2, no. 11, pp. 453, 454; see stars]
Littorina 529
“Littorina” incisa Yokoyama, 1927
(Pl. 358, figs. 4, 5)
Range—Pliocene of Japan.
Remarks—As pointed out by Habe (in litt.,
1971), this 5 mm. shell is a member of the
Pyramidellidae. We erroneously considered it to
be a Littorina (Littorinopsis) in our last number of
Indo-Pacific Mollusca, vol. 2, no. 11, p. 466 [p.
052340]. pl 358, fase4, Dp.
Synonymy—
1927 Littorina incisa Yokoyama, Journal of the Faculty of
Science, Imperial University of Tokyo, section I,
Geology, Mineralogy, Geography, Seismology, vol. 2,
part 4, p. 175, pl. 47, fig. 8 (Pliocene, Nagaya, Kaga,
Japan); holotype in Geological Institute, Imperial
University of Tokyo: 5 « 2.5 mm.
1970 Littorina incisa Yokoyama, Rosewater, Indo-Pacific
Mollusca, vol. 2, no. 11, p. 466.
Littorina kozaiensis Nomura and Onisi, 1940
(Pl. 349, figs. 6, 7)
Range—Lower Miocene of Japan.
Remarks—This species was described as re-
sembling L. adonis Yokoyama (see below), but as
having a larger number of spiral grooves. The
unique holotype (pl. 349, figs. 6, 7, a copy of the
original illustration) offers little basis for com-
parison with Recent species.
Synonymy—
1940 Littorina kozaiensis Nomura and Onisi, Japanese Jour-
nal of Geology and Geography, vol. 17, nos. 3 and 4,
p. 191, pl. 19, fig 6 a,b. (Y6suibori, Simizu, Kozaimura,
Japan); holotype: Saito H6-on Kai Museum, Register
No. 21762; 11 x 8 mm.
Littorina adonis Yokoyama, 1927
(Pl. 349, figs. 8, 9)
Range—Pliocene of Japan.
Remarks—Placed provisionally here in the
subgenus Littoraria, this species resembles L.
undulata, although the strong spiral sculpture is
also reminiscent of Littorinopsis, i.e. L. scabra,
etc. Unfortunately the outer lip and a portion of
the body whorl of the type (pl. 349, figs. 8, 9) are
missing and it is difficult to be sure of their exact
shape.
Synonymy—
1927 Littorina adonis Yokoyama, Journal of the Faculty of
Science Imperial University of Tokyo, section 2, vol.
1, part 10, p. 451, pl. 51, fig. 8, (Upper Musashino,
Koyasu southern Musashi, Japan); (unique holotype
in collection of Geological Institute Imperial Univer-
sity of Tokyo: 6 x 4 mm.).
Littorina lucida Yokoyama, 1927
(Pl. 349, figs. 4, 5)
Range—Pliocene of Japan.
Remarks—This species is from the same de-
posit as L. adonis but lacks the deeply incised
spiral sculpture. The type of lucida, although of
approximately the same size as adonis is more
slender.
We overlooked the fact that Habe, 1942 (Venus,
vol. 12, p. 37) and Abbott, 1958 (Proc. Acad. Nat.
Sci. Phila., vol. 110, p. 270) had pointed out that
L. lucida was a synonym of Assiminea japonica
von Martens, 1877.
Synonymy—
1927 Littorina lucida Yokoyama, Journal of the Faculty of
Science Imperial University of Tokyo, section 2, vol.
1, part 10, p. 451, pl. 51, fig. 9 (Upper Musashino,
Koyasu southern Musashi, Japan; unique holotype in
collection of Geological Institute Imperial University
of Tokyo: 5 * 3 mm.).
[05-317a]
530
Littoraria
Joseph Rosewater
Littorinidae
[ These occasional blank areas occur between
genera and subgenera to permit the insertion
of new material and future sections in their
proper systematic sequence.]
[05-318a]
January 15, 1972 INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
Index 531
INDEX TO LITTORINIDAE NAMES IN VOL. 2, NOS. 11 and 12
Looseleaf subscribers should place this index at the begin-
ning of the family Littorinidae, and just after the index tab
Littorinidae, p. 05-250.
The number following the name refers to the pagination
found at the top of the page. The column at the right is the
looseleaf pagination. All new names are in boldface type.
[looseleaf]
acuminata Gould, 438 05-300
acuta Menke, 469 05-353
acutispira E.A. Smith, 451 05-312
adelaidensis Cotton, 528 05-470
adonis Yokoyama, 453 05-317
afer Férussac, 431 05-279
africana Philippi, 476 05-360
albicans Metcalfe, 455 05-329
459 05-333
Algaroda Dall, 431 05-279
ambigua “Nuttall” Philippi, 448 05-310
ambiqua Weinkauff, 448 05-310
angulifera Lamarck, 455 05-329
antipodum Philippi, 471 05-355
arboricola Reeve, 459 05=333
ardouinianum Heude, 460 05-334
articulata “Menke” Philippi, 459 05-333
articulata Nevill, 464 05-338
articulata Philippi, 459 052333
australis Gray, 485 05-379
Austrolittorina Rosewater, 467 05-351
Bacalia H. & A. Adams, 431 05-279
Bacalia Gray, 431 05-279
balteata Reeve, 434 05-286
basterotii Payraudeau, 431 05-279
beccarii Tapparone-Canefri, 425 05-269
biangulata von Martens, 499 05-403
bicolor Lamarck, 519 05-455
blandfordi Dunker, 460 05-334
brevicula Philippi, 433 05-285
bullatus Martyn, 517 05-453
carinata Schrenck, 434 05-286
carinifera Menke, 464 05-338
carnifera Sherborn, 465 05-339
cecillei Philippi, 483 05-377
chaoi Yen, 493 05-397
Cidaris Réding, 513 05-449
cincta Quoy & Gaimard, 474 05-358
cinerea Pease, 505 05-409
cingulata Philippi, 459 05-333
coccinea Gmelin, 439 05-301
coccinea Martyn, 440 05-302
columna “Jonas” Philippi, 438 05-300
concolor Weinkauff, 438
460
conica Philippi, 465
contracta Nevill, 438
Corneolitorina Powell, 424
coronaria Lamarck, 517
coronatus Valenciennes, 515
crassior Philippi, 469
Cremnoconchus Blanford, 417
cumingi Philippi, 525
decollata Philippi, 477
delicatula Nevill, 460
diemensis Gray, 469
diemensis Quoy & Gaimard, 469
draparnauldii Audouin, 476
Echinella Swainson, 513
525
Echinellopsis Rovereto, 513
Echinininae Rosewater, 510
525
Echininus Clench & Abbott, 525
echinulatus Alth, 527 (Trochus)
echinulatus Kiener, 527
elegans Philippi, 515
erronea Nevill, 460
493
eudeli Sowerby, 493
exigua Dunker, 500
Ezolittorina Habe, 431
Fectaria Philippi, 513
feejeensis Reeve, 500
fernandezensis Rosewater, 471
fijiensis “Reeve” Nevill, 500
flammea Philippi, 459
flammulata Philippi, 459
foliorum Rumphius, 459
fortunei Reeve, 459
fragilis Fenaux, 426
glabrata Philippi, 445
globulus Angas, 426
gracilior Philippi, 443
grandinatus Gmelin, 516
granicostata E.A. Smith, 493
granocostata Reeve, 492
granularis “Gray” of authors, 492
502
Granulilittorina Habe & Kosuge, 491
guttata Reeve, 476
[05-257]
05-300
05-334
05-339
05-300
05-268
05-453
05-451
05-353
05-261
05-467
05-361
05-334
05-353
05-353
05-360
05-449
05-467
05-449
05-444
05-467
05-467
05-469
05-469
05-451
05-334
05-397
05-397
05-404
05-279
05-449
05-404
05-355
05-404
05-333
05-333
05-333
05-333
05-270
05-307
05-270
05-305
05-452
05-397
05-396
05-396
05-406
05-395
05-360
532 Index
Joseph Rosewater
hagruma Tokioka & Habe, 493
Hamus “Klein” H. & A. Adams, 513
heterospiralis Grabau & King, 434
hisseyiana Tenison-Woods, 426
hybrids, 488
incisa Yokoyama, 466
529
indistincta Turton, 479
infans E.A. Smith, 452
insularis E.A. Smith, 493
intermedia Philippi, 459
Isonema “Hall” Provancher, 431
iwakiana Nomura & Hatai, 506
knysnaensis Philippi, 478
kowiensis Turton, 479
kozaiensis Nomura & Onisi, 453
kraussi Rosewater, 444
laevigata Schrenck, 434
Laevilacunaria Powell, 424
Laevilitorina Pfeffer, 424
laevior Nevill, 465
laevis Philippi, 443
laevis “Philippi” Reeve, 469
Lamellilitorina Tryon, 455
lamellosa Montrouzier, 426
leucosticta Philippi, 497
leucosticta “Philippi” Tryon, 499
limax Gray, 440
lineatum Gmelin, 459
Litonia Weinkauff, 431
Litorina Menke, 431
Litorinna Dall, 431
Litormia Morch, 431
Littoraria Griffith & Pidgeon, 435
littorea Linné, 431
Littorelaea Leach, 431
Littorina Férussac, 431
Littorina-capsula, 483
493
Littorinopsis Mérch, 455
Littorivaga Dall, 431
Littornia Jenkins & Grocock, 431
Littorrina Gabb, 431
luchuana Pilsbry, 528
lucida Yokoyama, 453
luctuosa Reeve, 474
lutea Philippi, 459
luteola Quoy & Gaimard, 459
05-397
05-449
05-286
05-270
05-382
05-340
05-317a
05-363
05-314
05-397
05-333
05-279
05-410
05-362
05-363
05-317
05-306
05-286
05-268
05-268
05-339
05-305
05-353
05-329
05-270
05-401
05-403
05-302
05-333
05-279
05-279
05-279
05-279
05-297
05-279
05-279
05-279
05-377
05-397
05-329
05-279
05-279
05-279
05-470
05-317
05-358
05-333
05-333
Littorinidae
Macquariella Finlay, 424
major Swainson, 519
malaccana Philippi, 483
mandshurica Schrenck, 434
manschurica “Schrenck” Tryon, 434
marmorata Pfeiffer, 492
marmorata Philippi, 492
mauriciana “Lamarck” Potiez &
Michaud, 443
mauritiana Lamarck, 442
melanacme E.A. Smith, 493
melanostoma Gray, 462
miliaris Quoy & Gaimard, 492
millegrana Philippi, 491
492,
minor Weinkauff, 460
miodelicatula Oyama, 466
moerchi Adams & Angas, 426
monilifera Eydoux & Souleyet, 483
multistriata Tokioka, 483
natalensis Philippi, 489
Neritoides T. Brown, 431
Neritotrema “Recluz”” Wenz, 431
Neritrema Récluz, 431
newcombi Reeve, 459
newkombi “Reeve” in Weinkauff, 460
Nina Gray, 525
Nodilittorina von Martens, 481
nodosa Gray, 487
nodulosus Gmelin, 483
novaehiberniae Lesson, 459
novaezelandiae Reeve, 492
obesa Sowerby, 440
obtusata Linné, 431
oliveri Finlay, 471
opercula, 509
Pagodella Swainson, 513
Pagodus Gray, 513
pagodus Linné, 518
pallescens Philippi, 459
papillosa Lamarck, 521
papillosa of authors, 515
pascua Rosewater, 484
paytensis Philippi, 443
Pellilacunella Powell, 424
Pellilitorina Pfeffer, 425
penitaria Wood in G. Nevill, 472
perdix King & Broderip, 465
[05-258]
05-268
05-455
05-377
05-286
05-286
05-396
05-396
05-305
05-304
05-397
05-336
05-396
05-395
05-396
05-334
05-340
05-270
05-377
05-377
05-383
05-279
05-279
05-279
05-333
05-334
05-467
05-375
05-38 1
05-377
05-333
05-396
05-302
05-279
05-355
05-443
05-449
05-449
05-454
05-333
05-457
05-451
05-378
05-305
05-268
05-269
05-356
05-339
January 15, 1972
INDO-PACIFIC MOLLUSCA, vol. 2, no. 12
Index 533
perplexa Turton, 476
philippiana Habe & Kosuge, 493
philippiana Reeve, 459
philippina von Martens, 460
pica “Reeve” Turton, 479
picea Reeve, 479
picta Philippi, 503
pindata Philippi, 448
pintado Wood, 447
planaxis “Nuttall” Tryon, 504
praetermissa May, 445
Problitora Iredale, 426
pseudolaevis Nevill, 469
pulchella Dunker, 476
pultneyii Leach, 431
punctata Gmelin, 474
punctata Philippi, 459
pyramidalis Nevill, 465
pyramidalis Quoy & Gaimard, 481
483
quadriseriata Philippi, 515
radiata Eydoux & Souleyet, 492
rehderi Ladd, 524
reproduction of Tectarius, 510
reticulata Anton, 426
rhodea Biggs, 460
rietensis Turton, 479
Rissolittorina Ponder, 425
rubra Philippi, 459
rubropicta von Martens, 465
rugosa Menke, 487
rugosus Wood, 514
rusticus Philippi, 522
scabra Linné, 456
schmitti Bartsch & Rehder, 449
serialis Eydoux & Souleyet, 448
sieboldii Philippi, 459
sinensis Philippi, 459
sitchana Philippi, 431
songoense K. Martin, 523
souverbiana Crosse, 434
spinulosus Philippi, 527
05-360
05-397
05-333
05-334
05-363
05-363
05-407
05-310
05-309
05-408
05-307
05-270
05-353
05-360
05-279
05-358
05-333
05-339
05-375
05-377
05-451
05-396
05-460
05-444
05-270
05-334
05-363
05-269
05-333
05-339
05-381
05-450
05-458
05330
05-311
05-310
05-333
05-333
05-279
05-459
05-286
05-469
squalida Broderip & Sowerby, 431
strigata Lischke, 460
strigata Philippi, 459
strubelli von Martens, 493
subangulata Lamarck, 455
subcingulata Nevill, 460
Subditotectarius Ladd, 524
subgranosa Dunker, 499
subinermis Philippi, 523
subintermedia Nevill, 438
subnodosa Philippi, 495
suleatula Nevill, 438
sulcatus Nilsson, 431
sulculosa Philippi, 459
sundaica Altena, 450
suturalis Philippi, 459
syriaca Philippi, 476
Tectariinae Rosewater, 510
513
Tectarium, 523
Tectarius Valenciennes, 513
tectumpersicum Linné, 520
tenebrata “Nuttall” Jay, 448
tenuis Nevill, 460
tenuis Philippi, 438
trochiformis Dillwyn, 483
trochoides Gray, 483
tryphena Bartsch, 479
undulata Gray, 436
unifasciata Gray, 467
urieli Biggs, 493
ventricosa Philippi, 459
4992
vera Philippi, 521
verus J.E. Gray, 519
vidua Gould, 492
vilis “Menke” Philippi, 483
vitensis Dunker, 500
vitiensis “Reeve” von Martens, 500
vitrea Deshayes; 443
zebra Donovan, 435
zelandiae Finlay, 474
[05-259]
05-279
05-334
05-333
05-397
05-329
05-334
05-460
05-403
05-459
05-300
05-399
05-300
05-279
05-333
05-312
05-333
05-360
05-444
05-449
05-459
05-449
05-456
05-310
05-334
05-300
05-377
05-377
05-363
05-298
05-351
05-397
05-333
05-396
05-457
05-455
05-396
05-377
05-404
05-404
05-305
05-297
05-358
534 Index
Joseph Rosewater
Littorinidae
Published by
THE DEPARTMENT OF MOLLUSKS
Delaware Museum of Natural History
Box 3937, Greenville, Delaware
19807, U.S.A.
[05-258]
——
INDO-PACIFIC
MOLLUSCA
Monographs of the Marine Mollusks of the World with Emphasis
on those of the Tropical Western Pacific and Indian Oceans
EDITED BY
R. TUCKER ABBOTT
VOLUME 3
Published by
DELAWARE MUSEUM OF NATURAL HISTORY
Box 3937, Greenville,
Delaware 19807, U.S.A.
74 Title Page
Editors
Volume 3
Notice
Looseleaf subscribers should place this title page in the first
binder immediately after the title page of volume 2. It will
be your record of the number of volumes in your looseleaf set.
[this sheet issued June 1, 1973, with vol. 3, no. 14]
May 30, 1973
M\oll
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13
Drupa 1
WILLIAM H. DALL
SECTIONAL LIBRARY
DIVISION OF MOLLUSKS
THE GENUS DRUPA IN THE INDO-PACIFIC
by WituiaAM K. EMERSON
Department of Living Invertebrates
The American Museum of Natural History
New York, N.Y. 10024
and
WALTER O. CERNOHORSKY
Auckland Institute and Museum
Auckland, New Zealand
Abstract
A revised classification of the gastropod
genus Drupa Roding (Muricidae: Thaidinae) is
presented. The following taxa are recognized:
Drupa (Drupa) morum morum Roding, 1798;
D. (D.) morum iodostoma (Lesson, 1840); D.
(D.) ricinus ricinus (Linnaeus, 1758); D. (D.)
ricinus hadari Emerson and Cernohorsky, new
subspecies; D. (D.) elegans (Broderip and
Sowerby, 1829); D. (Ricinella) rubusidaeus
Roding, 1798; D. (R.) speciosa (Dunker, 1867);
D. (R.) clathrata clathrata| (Lamarck, 1816);
D. (R.) clathrata miticula (Lamarck, 1822); D.
(Drupina) grossularia Roding, 1798; and D.
(Drupina) lobata (Blainville, 1832). Generic
and specific synonymies are given for these
taxa, together with distributional and ecolog-
ical data for each species.
Indo-Pacific Drupa
Species of the genus Drupa Roding are con-
fined in their distribution to the tropical Indo-
Pacific region, where they are commonly en-
countered on_ intertidal reef-flats. Drupa
species are muricacean’ gastropods which
show a close relationship with the larger, but
otherwise similar and closely related species
of Thais Roding, and are currently assigned to
the subfamily Thaidinae within the family
Muricidae. Because of the close similarity in
shell morphology, species of Drupa Réding,
and Morula Schumacher have frequently been
considered to be congeneric or only subgener-
ically separable. Wu (1965b), in his compara-
tive study of the functional anatomy of the
digestive systems of Drupa ricinus (Linnaeus)
and Morula granulata (Duclos), found mor-
phological differences in features of the rad-
ula, gland-gut complex, stomach and _ rectal
gland, and most notably in the structure of the
salivary glands.
The exterior of the animal of Drupa consists
of a foot with an attached, chitinous oper-
culum, a head, snout, proboscis, a pair of ten-
tacles and eyes and the reproductive organ.
Animals are dioecious, with the male’s penis
situated behind the right tentacle below the
thin mantle. The operculum is brown in color
and corneous, stereotyped thaidine in appear-
oe ae me Pe
iS SOP . a OF Sad
Plate 1. Camouflage in Drupa (Drupina) grossularia Réding.
Cape Tuiolemu, Upolu, Samoa Ids. Top figure: Two
specimens in situ on an exposed algae-covered reef. Bot-
tom figure: two over-turned, living specimens on _ reef
(photo courtesy A. Solem).
[14-801]
2 Emerson and Cernohorsky Muricidae
ance, with a series of plateaulike ridges on the occupying the mid-eulittoral zone. Demond
side of attachment and concentric growth-
rings on the exterior which converge basally
into an ill-defined nucleus (see pl. 3, _ figs.
1-4).
In the Fiji Islands and the New Hebrides,
Drupa were encountered in the mid-eulittoral
and upper eulittoral region of the intertidal
zone, generally on windward and_ exposed
reef-flats. Species were considerably less nu-
merous on protected, coral-strewn or shingle-
covered leeward reefs. Drupa morum Réding,
D. rubusidaeus Réding, and D. ricinus (Lin-
naeus) were most frequently collected on ex-
posed algal ridges which were constantly kept
moist through agitated waters and a_ breaking
surf. The same species occurred on reef-edges
covered with calcareous algae and_ detritus.
Drupa ricinus, however, also occurred in the
mid-eulittoral zone, while D. — grossularia
Roéding, was usually confined to this part of
the reef-zone. Kay (1971), in her study of the
molluscan fauna of Fanning Island in the Line
Islands, reported D. ricinus, D. morum and D.
grossularia to be the most common macro-
mollusks inhabiting reef-flats. D. ricinus and
D. grossularia were among the dominant spe-
cies of the beach-rock assemblages, with the
former species restricted to exposed areas.
D. ricinus also occurred on a_ subtidal, la-
goonal patch reef, but was found to be far
more abundant on seaward reefs than on the
patch reefs in the lagoon. Salvat (1970), in his
study of littoral molluscs of Fangataufa, Tua-
motus, found a_ similar distributional pattern
of Drupa as that observed by the junior author
in Melanesia, with D. ricinus being more fre-
quent on seaward algal ridges and D. morum
Figs. 1-3. Drupa (Drupa) morum morum Réding, 1798.
1, Chisimaio, Somalia (ANSP 298192); 2, 3, adult and
immature, both from Okinawa Id. (ANSP 302877).
Figs. 4, 5. Drupa (Drupa) morum iodostoma (Lesson, 1840).
Both from Ua Huka Id., Marquesas Ids. (ANSP 155617
and 156169).
Figs. 6-8, 11. Drupa (Drupa) ricinus ricinus (Linnaeus, 1758).
6. Okinawa Id. (ANSP 302919); 7, 11, Mahé. Seychelles
Ids. (ANSP 266229): 8, immature, Moorea, Society Ids.
(ANSP 283222).
Figs. 9, 10. Drupa (Drupa) ricinus new subspecies hadari
Emerson and Cernohorsky. 9, paratype (AMNH 112617a);
10, holotype (AMNHII 166928). Eilat, Gulf of Aqaba, Israel.
Fig. 12. Drupa (Drupa) elegans (Broderip and Sowerby,
1829). Society Islands (ANSP 199558).
Figs. 13-15. Drupa (Ricinella) rubusidaeus Réding, 1798.
13, Gesira, Somalia (ANSP 299187); 14, Isles Radama,
N.W. Madagascar (ANSP 257243); 15, immature; Malaita
Id., British Solomon Ids. (ANSP 289624).
(1957) reported D. grossularia as occurring in
Micronesia most commonly on windward reef
flats on rocks near the low tide line, less fre-
quently on leeward reefs and rarely in la-
goons. D. morum morum was recorded com-
monly found living among rocks and coral of
windward reef-flats and on windward _la-
goonal reef-flats of the larger atolls, but rarely
on leeward reefs. This species was encoun-
tered most often near the low tide line, on or
near the reef-edge, but also was taken in tide
pools across the entire reef-flat. D. ricinus was
reported to occupy a similar habitat to D.
morum morum, while D. rubusidaeus was
found to inhabit both windward and leeward
reef-flats, living under rocks and coral rubble;
it also occurred in tide-pools near, or just be-
low the low tide line, and was found also liv-
ing on coral heads off the seward reef-edge, in
10 to 15 feet of water. Heinicke (1970) en-
countered D. lobata (Blainville), seemingly al-
ways occurring in pairs, in the lagoonal chan-
nel among coral heads at Diani Beach, Kenya.
Drupa species are commonly encrusted with
algae, coral growth, vermetids, Foraminifera,
Hipponix and other’ extraneous’ organisms,
making them blend in with the substrate upon
which they rest (see pl. 1). Nothing is known
about natural enemies of Drupa, except that
Schoenberg (1971) records captive Conus tex-
tile Linnaeus preying on Drupa morum, “D.
speciosa” (= D. rubusidaeus), and both color
forms of D. ricinus, among numerous other
species of Hawaiian prosobranch gastropods.
Conflicting reports may be found in litera-
ture on the feeding habits of Drupa species.
Salvat (1970) examined the microscopic con-
Figs. 16-18. Drupa (Ricinella) clathrata clathrata (Lamarck,
1816). 16, Okinawa Id. (ANSP 289725); 17, Hivaoa Id.,
Marquesa Ids. (ANSP 155492); 18, Pacific Ocean (ANSP
36720).
Figs. 19, 20. Drupa (Ricinella) clathrata miticula (Lamarck,
1822). 19, Arsenal Bay, Mauritius (ANSP 273087); 20,
Mahébourg, Mauritius (AMNH 104995).
Figs. 21, 22. Drupa (Ricinella) speciosa (Dunker, 1867). Both
from “Rarotonga, Cook Ids.’—probably an error (ANSP
29873). Known from the Tuamotus and Pitcairn Islands.
Figs. 23, 24. Drupa (Drupina) grossularia Roding, 1798.
23, Okinawa Id. (ANSP 225428); 24, immature, Sorsogon.
Luzon Id., Philippines (ANSP 224148).
Figs. 25, 26. Drupa (Drupina) lobata (Blainville, 1832). 25,
Mogadiscio, Somalia (ANSP 295772); 26, Direction Id.,
Cocos-Keeling Ids., Indian Ocean (ANSP 288455).
(all figures about natural size)
[14-802]
May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 3
Plate 2. Genus Drupa Roding in the Indo-Pacific
(all figures about natural size)
Explanation on opposite page.
[14-803]
4 Emerson and Cernohorsky
tents of the digestive tracts of several reef-
dwelling gastropods and decided that D. rici-
nus and D. morum morum were herbivores,
whereas other authors (Wu 1965b; Taylor
1968; Kay 1971; Cemohorsky, personal obser-
vation) report these species to be carnivores.
The feeding habits of the following species
have been reported:
Drupa morum morum
worms and sipunculids (Kay, 1971) [Line
Ids. ]
barnacle _— Tetraclita
1968) [Seychelles Ids. ]
herbivorous (Salvat, 1970) [Tuamotu Ids. }
sipunculid worms (Cernohorsky, pers. ob-
servation) [New Hebrides]
Drupa ricinus
live prey, i.e. sponges and_holothurians,
or carrion (Wu, 1965b) [Hawaiian Ids. ]
molluscs, barnacles and worms’ (Kay
1971) [Line Ids.]
Drupa grossularia
omnivorous (Salvat, 1970) [Tuamotu Ids. ]
squamosa (Taylor,
Plate 3. Opercula of Drupa.
Figs. 1, 2. Drupa (Drupa) morum morum Réding, from
Nananu-i-Ra Id., Fiji Ids. 1, inner surface; 2, outer
surface.
Fig. 3. Drupa (Ricinella) clathrata clathrata (Lamarck),
from Pango Point, Efate Id., New Hebrides.
Fig. 4. Drupa (Drupina) grossularia Réding, from Wadigi
Id., Fiji Ids.
Muricidae
In comparison to Drupa_ species, Morula
granulata (Duclos) was reported to feed on
other mollusks and bamacles in the Seychelles
Islands (Taylor, 1968), but was recorded prey-
ing on holothurians, boring into bivalves of
Isognomon and Ostrea, and consuming carrion
in the Hawaiian Islands (Wu, 1965b).
No information is recorded on the mode of
reproduction of Drupa, although J. B. Taylor
(in litt.) reports certain Hawaiian species of
Drupa (sensu lato) to have planktonic veli-
gers. Such a larval stage would account for
the wide distribution of most species.
Radulae
The radula of Drupa is of the rachiglossate
type, with 3 teeth per transverse row with a
formula of 1-1-1. The radular ribbon is small
and very narrow, and the lateral teeth are
simple and sickle-shaped (see pl. 4, fig. B).
The rachidian teeth are more or less subquad-
rate or rectangular, the base is weakly con-
cave, the central cusp is slender and longer
than the flanking, bifid to quadrifid side-
cusps. The lateral denticles are small, moder-
ately deeply rooted and number from 2 to 5,
and the end-cusps are usually slightly larger
than the lateral denticles (see pl. 4, fig. A).
Although the shells of Drupa show an af-
finity with species of Morula Schumacher, the
radulae of the moruline group of species dif-
fer in the following particulars: the central
cusp of the rachidian is more deeply rooted,
the flanking side-cusps are not multifid as in
Drupa, but are simple, and the central cusp
and side-cusps are separated from each other
by an interposing small, intermediate cusp.
The radula of Morula is essentially muricine
in appearance while that of Drupa is a weakly
modified thaidine radula, which approaches
that of Murex s. s.
A classification based on radular morphol-
ogy is complicated by the sporadic appear-
ance of a drupine-type radula in other thai-
dine genera, e.g. Agnewia tritoniformis (Blain-
ville, 1832) [see Kesteven, 1902, pl. 29, fig. 5
and Cooke, 1919, text fig. 26], Semiricinula
muricina (Blainville, 1832) [see Arakawa,
1965, pl. 14, figs. 19, 20], and Neothais smithi
(Brazier, 1889). Wu (1965b) surmized that the
distinctive drupine and moruline radula_pat-
tern displayed by the species investigated by
him may be directly associated with their re-
spective feeding habits.
[14-804]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13
Drupa 5
A
Plate 4. Radula of Drupa (Drupa) ricinus ricinus (Linnaeus).
One transverse row. Fig. A. rachidian or central tooth.
Fig. B. lateral tooth.
Abbreviations—
ec, end cusps
Id, lateral denticles
sc, side cusps
cc, central cusp
Distribution and fossil record
Species of Drupa are largely confined in
distribution to the Indo-Pacific region, but
some range from the northern regions of the
Red Sea to Easter, Clipperton and the Gala~
pagos Islands in the eastern Pacific Ocean.
They inhabit tropical waters and do not occur
beyond latitudes of 35°N and 35°S. Although
muricacean gastropods date from the _ late
Mesozoic in the Cretaceous, typical thaid gas-
tropods first appear in the Oligocene in mid-
Tertiary (Keen, 1971). These Cenozoic forms
apparently are the precursors of the drupine
forms that are here referred to Drupa s.l. The
drupine forms, with low-spired, — thaidlike
shells having strong labial and columellar
teeth, are poorly represented in the fossil
record and are known only from the Pleisto-
cene. Fossil moruline forms, with smaller,
oval-biconical and higher-spired_ shells than
those of Drupa s.l., are recorded as ranging
from the Eocene to the Pleistocene (Eames,
1971, as “Drupa (Sistrum)”). The limited pale-
ontological data suggest, therefore, that Drupa
sl. and Morula_ s.l. evolved independently
from a pre-thaid stock. The fossil evidence is
not sufficient, however, to establish a_ well-
documented evolutionary chronology of these
generic groups.
As pointed out by Hertlein (1960), Paleogene
fossils from Europe and the East Indies have
been placed in the genera “Ricinula” or “Sis-
trum.” A survey of the literature indicates that
some of these taxa are not closely allied to
Drupa or Morula. Cossmann (1889, p. 132;
1903, p. 80, pl. 3, fig. 15) refers Purpura
ringens Deshayes, 1865, from the Eocene of
the Paris Basin to Ricinula, a spurious drupine
assignment. Another example is Sistrum bay-
lei Cossmann & Lambert, (1884, p. 175, pl. 5,
fig. 19), from the Oligocene of France; both of
these taxa appear to be buccinacean gastro-
pods. Purpura styriaca Stur in Hilber, 1879, a
Neogene fossil from the Miocene of Hungary,
is a thaid, although this taxon was recently
referred to Drupa by Strausz (1966, p. 284, fig.
130).
Ricinula puruensis Martin (1914, p. 147, pl.
4, fig. 104) [not fig. 105], described from the
late Eocene of Java, appears to be the earliest
record for the genus Morula. Miocene species
of Morula s.l. include: M. angsanana (Martin,
1921, p. 466, pl. 59, fig. 52), M. turrita (Martin,
1880, p. 41, pl. 8, fig. 3), both from Java, M.
austriaca (Hoernes & Auinger, 1882, pl. 16,
figs. 14-17) from the Vienna Basin, and M. in-
constans (Michelotti, 1847, p. 217) from Italy.
Neogene fossils that are purported to repre-
sent living species of Morula s.l. are reported
from Mio-Pliocene, Pliocene and_ Pleistocene
deposits in the’ present Indo-Pacific faunal
region and elsewhere.
Drupa s.str. is recorded by Eames (1971) as
ranging from the Pliocene to Recent and as
occurring in the Red Sea, Indo-Pacific, East
Africa) and questionably in North America.
The New World record is based on the genus
Condonia Hertlein, 1965, (type-species by orig-
inal designation Sistrum hannai Howe, 1922),
from the Pliocene of Oregon and California.
This species, however, bears a resemblance
to certain buccinacean shells, such as Can-
tharus and Columbella, and the monotypic
genus Condonia, therefore appears to be ref-
erable to the superfamily Buccinacea instead
of the Muricacea.
Pentadactylus rhombiformis Martin (1899,
p. 138, pl. 21, figs. 316a, b), described from
the Pliocene of Java, was compared with sev-
eral living muricacean species, including ref-
erences to Reeve’s (1846, Purpura, pl. 3, fig.
13) illustration of Drupa rubusidaeus Roding.
The species was considered closest in relation-
ship to Purpura muricina Blainville, and the
description and illustration of the Indonesian
fossil suggest that it is a spinose thaid.
Thus the available data indicate no valid
records for Drupa s.l. prior to the Pleistocene.
The following species are reported from Pleis-
tocene deposits: Drupa (Drupa) morum
Roding, 1798; D. (D.) ricinus ricinus (Linnaeus,
[14-805]
1758), and Drupa (Ricinella) rubusidaeus Réd-
ing, 1798 (see distributional records).
Classification
The Linnaean species of Drupa_ s.l. were
originally described in the genus Murex Lin-
naeus. Rodding (1798) proposed the genus
Drupa; Montfort (1810) the genus Sistrum;
and Lamarck (1816) the genus Ricinula, all
for some of the drupine species previously
assigned to Murex. The genus-group name
Ricinula remained in use in malacological lit-
erature until about 1913, when it was grad-
ually replaced with Réding’s chronologically
prior Drupa.
Thiele (1929) accepted Drupa as a_ valid
genus-group, but relegated several moruline
genera, i.e. Cronia H. & A. Adams, Morulina
Dall (=Azumamorula’ Emerson), — Phrygio-
murex Dall, Maculotriton Dall and Drupella
Thiele, as subgenera of Drupa. Wenz (1941)
erected the new subfamily Drupinae, which
besides the type-genus Drupa, contained the
genus Thais R6ding and other thaid genera,
together with the non-thaid genus Tritonalia
Fleming (= Ocenebra Gray). Drupinae Wenz,
1938 and 1941 is presently considered a junior
synonym of Thaidinae Suter, 1909 (as Thai-
sidae Suter, 1909, Rec. Canterbury Mus., vol.
1, p. 11; 1909, Subantarctic Islands of New
Zealand, art. 1, p. 27). The subfamilial name
Thaidinae was conserved by action of the
International Commission on Zoological No-
menclature (Opinion 886, 1969) in preference
to the chronologically older, but less fre-
quently used Purpurinae.
Species of Drupa are here assigned to the
genus mainly on_ shell-morphology, although
radular characters have also been considered.
As pointed out in the section on “Radulae,” a
classification on radular characters alone
would require an inclusion of species of Ag-
newia, Semiricinula, Neothais and a species of
Morula. Such a classification was in fact pro-
posed by Cooke (1919), who included 7 non-
drupine species in Drupa on the basis of
radular characters which he considered “dis-
tinctly of the Drupa type.” It is obvious that
on shell-morphology alone, the limits of Drupa
are well-defined, but radular characters of
Drupa also appear rarely in species refer-
able to other thaidine genera.
Due to the dispersal at auction of the Bolten
collection, on whose specimens Réding’s new
Emerson and Cernohorsky
Muricidae
descriptions were based, the whereabouts of
the type-specimens are no longer known. F. C.
Schmidt did purchase a small part of Bolten’s
collection in 1819, but in a letter written by
him, he observed that a great amount of the
collection was purchased by Hamburg buyers.
From those specimens procured by Schmidt
from the Bolten collection, now in the Natur-
kundemuseum, Staatliche Museen zu _ Gotha,
Germany, only very few can be traced back to
Bolten (Dr. Motschmann, in litt.). In the ab-
sence of Rd6ding’s type-specimens we have
designated appropriate cited illustrations of
specimens figured by other authors as lecto-
types of Roding’s species.
List of Recognized Taxa
Below are listed the recognized generic and
specific taxa for the genus Drupa. The eleven
species and subspecies are referred to three
genus-groups. All are living, and three are
also recorded as Pleistocene fossils.
Family Muricidae Rafinesque, 1815
Subfamily Thaidinae Suter, 1909
Genus Drupa Réding, 1798
Subgenus Drupa Réding, 1798
morum morum Roding, 1798. Type species.
Recent. Indo-Pacific. and Eastern Pacific,
except the Marquesas Islands. Pleistocene.
morum iodostoma (Lesson, 1840). Recent,
Marquesas Islands.
ricinus ricinus (Linnaeus, 1758). Recent,
Indo-Pacific and Eastern Pacific. Pleisto-
cene.
ricinus hadari Emerson and _ Cernohorsky,
new subspecies. Recent, Red Sea.
elegans (Broderip and Sowerby, 1829). Re-
cent, Wake Island to the Tuamotu Islands.
Subgenus Ricinella Schumacher, 1817
rubusidaeus R6ding, 1798. Type species. Re-
cent, Indo-Pacific. Pleistocene.
speciosa (Dunker, 1867). Recent, Tuamotu
and Pitcairn Islands.
clathrata clathrata (Lamarck, 1816). Recent,
tropical west Pacific Ocean.
clathrata miticula (Lamarck, 1822). Recent,
Indian Ocean.
Subgenus Drupina Dall, 1923
grossularia Roding, 1798. Type species. Re-
cent, East Indian Ocean and Pacific.
lobata_ (Blainville, 1832). Recent,
Ocean.
Indian
[14-806]
May 30, 1973
Abbreviations
The following institutional abbreviations are
used in this paper:
AIM—Auckland Institute and Museum, Auck-
land
AMNH—American Museum of Natural History,
New York
AMS— Australian Museum, Sydney
ANSP—Academy of Natural Sciences of Phil-
adelphia
BM (NH)—British Museum (Natural History),
London
BPBM— Bernice P. Bishop Museum, Honolulu
DM—Dominion Museum, Wellington
DMNH—Delaware Museum of Natural History,
Greenville
FMNH—Field Museum of Natural History, Chi-
cago
MCZ—Museum of Comparative Zoology, Cam-
bridge, Massachusetts
LACMNH—Los Angeles County Museum of
Natural History
MHNG—Museum d Histoire Naturelle, Geneva
SDMNH—San Diego Museum of Natural His-
tory
USNM—National Museum of Natural History,
Washington, D.C.
WAM—Western Australian Museum, Perth
Acknowledgments
We gratefully acknowledge the help ex-
tended to us in providing access to collections,
technical assistance, loan of specimens, field
data and information on types. We would like
to thank the following persons:
R. T. Abbott—DMNH; E. Binder—MHNG;
W. J. Clench and R. D. Turner—MCZ; S. P.
Dance, National Museum of Wales, Cardiff;
R. K. Dell—DM; the late A. Hadar, Tel Aviv,
Israel; E. A. Kay, University of Hawaii, Hono-
lulu; J. Knudsen, University Zoological Mu-
seum, Copenhagen; Y. Kondo—BPBM,; Fei-Jann
Lin, Academia Sinica, Taipei, Taiwan; J. H.
McLean—LACMNH,; D. F. McMichael—formerly
AMS; D. Motschmann, Naturkundemuseum,
Gotha; W. E. Old, Jr.—AMNH; V. Orr Maes—
ANSP; A. W. B. Powell—AIM; H. A. Rehder,
J. Rosewater and J. P. E. Morrison—USNM;
G. E. Radwin—SDMNH; Mme. P. Revercé, Nou-
mea, New Caledonia; M. G. Richards—formerly
AMNH:; B. Salvat, Muséum National d’Histoire
Naturelle, Paris; V. Siewersten, Koloa, Hawaii;
A. Solem—FMNH; Mr. & Mrs. G. D. Stout,
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13
Drupa 7
New York; J. B. Taylor, Prescott College, Ari-
zona; J. Taylor and K. Way—BM (NH); N. Teb-
ble, Oxford University Museum; J. J. Wage-
man, Koloa, Hawaii; C. S. Weaver, Kailua,
Hawaii; B. R. Wilson and S. M. Slack-Smith—
WAM; Shi-Kuei Wu, University of Michigan,
Ann Arbor.
Species excluded from Drupa
Included under this heading are species of
Thaidinae which were originally described in
Drupa, or have been referred to this genus by
subsequent authors. In the latter category,
only those species requiring further explana-
tion have been listed.
Neothais bollonsi (Suter, 1906)
(Pls. 5, 6)
Remarks—Suter (1909) assigned this mor-
uline species to the genus Drupa on the basis
of the typically drupine radula. Iredale (1915)
correctly synonymized D. bollonsi with the
southeast Australian species Purpura smithi
Brazier, 1889, and assigned it to the genus
Neothais Iredale, 1912. Despite its drupine
radular characters, the species should be
placed near Morula Schumacher.
amas aan
/ Md hat i
Olam
Plate 5. Radula of Neothais smithi (Brazier). Half a trans-
verse row; Sunday Id., Kermadec Ids. [synonym is Drupa
bollonsi Suter].
Plate 6. Neothais smithi (Brazier). [synonym is Drupa_ bol-
lonsi Suter].
Fig. 1. Sunday Id., Kermadec Ids. (AIM; 24.5 x 17.4 mm).
Fig. 2. Norfolk Id. (AIM; 18.0 x 12.0 mm).
[14-807]
8 Emerson and Cernohorsky
Synonymy—
1889 Purpura smithi Brazier, Australian Museum Memoir,
no. 2, p. 28, pl. 4, figs. 1-4, 7-12, 21-22 (Lord Howe
Id.) [as Purpura (Polytropa) smithi on plate ex-
planation].
1902 Purpura tritoniformis var. smithi Brazier, Kesteven,
Proceedings of the Linnaean Society of New South
Wales, pt. 4, p. 534.
1906 Purpura striata Martyn subsp. bollonsi Suter, Trans-
actions and Proceedings New Zealand Institute,
vol. 38, p. 331 (Kermadec Ids.).
1909 Drupa bollonsi Suter, Proceedings of the Malacolog-
ical Society of London, vol. 8, p. 254, pl. 11, figs.
5-7 (shell, operculum and_ radula); 1913 Suter,
Manual of the New Zealand Mollusca, p. 428, pl.
19, fig. 11 (New Zealand).
1915 Neothais smithi (Brazier), Iredale, Transactions and
Proceedings New Zealand Institute, vol. 47, pp.
474, 475; 1915 Oliver, Transactions and Proceed-
ings New Zealand Institute, vol. 47, p. 536; 1950
Dell, Dominion Museum Records, Zoology, vol. 1,
no. 3, p. 26.
Condonia hannai (Howe, 1922)
Remarks—This moderately large, (66.8 mm
long) species from the Pliocene of Oregon and
California’ was originally described in the
genus Sistrum Montfort. Hertlein (1965) pro-
posed for this species the new genus Con-
donia, and placed the genus in the muricid
subfamily “Drupinae” with apparent reluc-
tance. He concluded that: “Condonia hannai
bears a general resemblance to the Recent
Drupa iodostoma Lesson but the spire of
that species is low and the columella [=error
for inner margin of outer lip] bears denticles
typical of Drupa.” He also noted that the
genus was not known as a fossil in the Eastern
Pacific region, but that living representatives
occur in the Galapagos Islands and at Clipper-
ton Island. Although the labial dentition does
superficially resemble that of Drupa, the type-
species of Condonia lacks columellar den-
ticles. This extinct species appears to be a buc-
cinacean gastropod, and Dr. G. E. Radwin
(in litt.) considers Condonia hannai to be an
extralimital representative of the genus Col-
umbella s.str. which, perhaps due to its un-
usual northern, cooler-water habitat, attained
a giant size. The monotypic genus Condonia,
therefore, appears to be referable to the Buc-
cinacea rather than the Muricacea.
Synonymy—
1922 Sistrum hannai Howe, Univ. California Publ., Bull.
Dept. Geol. Sci., vol. 14, no. 3, p. 102, pl. 8, figs.
1, 5 (Fossil Point, S. W. Empire City, Coos Bay, Plio-
cene of Oregon); 1943 Weaver, Univ. Washington
Publ. Geology, vol. 5, pt. 2, p. 450, pl. 87, figs. 14,
Muricidae
16 (figured holotype); 1960 Hertlein, Veliger, vol. 3,
no. 1, p. 8 (San Benito County, Pliocene of Cali-
fornia).
1965 Condonia hannai (Howe), Hertlein, Occas. Papers
California Acad. Sciences no. 49, p. 4, figs. 3, 4
(figured holotype).
Azumamorula mutica (Lamarck, 1816)
(Pls. 7, 8)
Remarks—For the western Indian Ocean spe-
cies Ricinula mutica Lamarck, which is inter-
mediate in shell-characters between Drupa
Roding, and Morula Schumacher, Dall (1923)
proposed the genus-group Morulina. Thiele
(1929) and Wenz (1941) assigned Morulina as
a subgenus to Drupa. Emerson (1968) pro-
posed the substitute name Azumamorula for
the preoccupied Morulina Dall (non Morulina
Borner, 1906, in Insecta), and figured the rad-
ula of the type-species, which is typically moru-
line.
Plate 7. Radula of Azumamorula mutica (Lamarck). Half a
transverse row; Black River Bay, Mauritius (after Azuma
and d’Attilio in Emerson, 1968).
Plate 8. Azumamorula mutica (Lamarck).
Fig. 1. Holotype from unknown locality (“Mozambique”
on label) [MHNG no. 1101/19; 20.2 x 16.5 mm].
Synonymy—
1816 Ricinula mutica Lamarck, Tabl. Encycl. Methodique,
p. 1, pl. 395, figs. 2a, b (no locality given); 1846
Reeve, Conchologia Iconica, vol. 3, pl. 2, fig. 1.
1822 Ricinula pisolina Lamarck, Hist. nat. anim. s. ver-
tebres, vol. 7, p. 233 (Ile de France - Mauritius);
1835 Kiener, Spec. gen. icon. coquilles vivantes,
vol. 8, p. 20, pl. 4, fig. 8a (juvenile specimen);
1844. Deshayes and Milne-Edwards, Hist. Nat.
anim. s. vertebres, ed. 2, vol. 10, p. 52.
1919 Morula mutica (Lamarck), Cooke, Proceedings of
the Malacological Society of London, vol. 13, p.
106 (description of radula).
[14-808]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 9
1923 Morulina mutica Lamarck, Dall, Proceedings of the
Academy of Natural Sciences of Philadelphia, vol.
75, p. 303.
1929 Drupa (Morulina) mutica (Lamarck), Thiele, Handb.
syst. Weichtierkunde, vol. 1, p. 294; 1941 Wenz,
Handb. Palaozoologie, vol. 6, pt. 5, p. 1112, text
fig. 3159.
1968 Azumamorula mutica ‘Lamarck), Emerson, Nautilus,
vol. 81, no. 4, p. 125, text fig. (radula).
Drupa vitiensis Pilsbry in
Pilsbry and Bryan, 1918
Remarks—The species illustrated by Pilsbry
and Bryan is the male form of Drupella cornus
(Roding, 1798). Generally credited to Pilsbry,
1921, the specific name was validated by Pils-
bry and Bryan through a published illustration
in combination with a specific name.
Synonymy—
1918 Drupa vitiensis Pilsbry in Pilsbry & Bryan, Nautilus,
vol. 31, no. 3, pl. 9, fig. 5.
1921 Sistrum’ vitiense Pilsbry, Proceedings of the Acad-
emy of Natural Sciences of Philadelphia, vol. 72,
p. 319 (Fiji Ids.).
Drupa walkerae Pilsbry and Bryan, 1918
Remarks—The species described by the au-
thors belongs to Drupella Thiele, 1925, and is
similar to, if not conspecific with D. rugosa
(Born, 1778).
Synonymy—
1918 Drupa walkerae Pilsbry & Bryan, Nautilus, vol. 31,
no. 3, p. 99, pl. 9, fig. 4 (Honolulu Harbor, Hawai-
ian Ids.).
Roding (1798) described in the Museum Bol-
tenianum, pp. 55-56, the following Drupa spe-
cies: D. chamaemorus, D. botroides, D. uva,
D. cornus, D. glans, D. muricina, D. mancinella,
D. trapa and D. aesculus. All these taxa are non-
drupine species referable to various thaidine
genera, with the exception of D. glans, which
belongs to the genus Austrofusus Kobelt, 1879,
in the family Buccinidae.
[14-809]
10 Emerson and Cernohorsky
Muricidae
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Mollusca Testacea. Mémoires de l'Institut d’Egypte, vol.
37, pp. 1-89.
Langdon, A. W. 1875. Shells of Ceylon. Quarterly Journal
of Conchology, vol. 1, pp. 71-78.
Mansfield, W. C. in H. T. Stearns and K. N. Vaksvik.
1935. Geology and Ground-Water resources of the Is-
land of Oahu, Hawaii. Territory of Hawaii, Division of
Hydrography Bulletin, 1, pp. 166-168.
Martin, K. 1880. Die Tertiarschichten auf Java. Leiden,
pp. 1-164, pls. 1-28, ] map.
Martin, K. 1899. Die Fossilien von Java. Mollusken. Parts
5-7. Samml. geol. Reichs-Museums Leiden, N.F., vol. 1,
Abt. 1, Heft 6-8, pp. 133-221, pls. 21-33.
Martin, K. 1914. Die Fauna des Obereocans von Nang-
gulan, auf Java. A. Gastropoda. Samml. geol. Reichs-
Museums Leiden, N.F., vol. 2, Abt. 2, Heft 5, pp. 107-
178, pls. 1-6.
Martin, K. 1921. Die Fossilien von Java. Die Mollusken
der Nijalindungschichten. Samml. geol. Reichs-Museums
Leiden, N.F., vol. 1, Abt. 2, Heft 3-4, pp. 446-496, pls.
58-71.
Melvill, J. C. 1909. Report on the marine Mollusca ob-
tained by Mr. J. Stanley Gardiner, F.R.S. among the
islands of the Indian Ocean in 1905. Transactions of the
Linnean Society of London, ser. 2, Zoology, vol. 13, pt.
1, pp. 65-138.
Melvill, J. C. and E. R. Sykes, 1899. Notes on a third collec-
tion of marine shells from the Andaman Islands, with
description of three new species of Mitra. Proceedings
of the Malacological Society of London, vol. 3, pp.
22()-229.
Melvill, J. C. and R. Standen. 1898. Report on the marine
Mollusca obtained in Torres Strait. Journal of the Lin-
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Michelotti, G. 1847. Description des fossiles des Terrains
Miocenes de IJ'Italie septentrionale. | Natuurkundige
Varhandelingen Hollandsche Maatschappij der Weten-
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Ostergaard, J. M. 1928. Fossil marine Molluscs of Oahu.
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Reeve, L. A. 1846. Conchologia Iconica. Monograph of the
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Index.
Salvat, B. 1970. Etudes quantitatives (comptages et bio-
masses) sur les Mollusques recifaux de TAtoll de
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no. 14, pp. 1-58, pls. 1-5.
Satyamurati, S. T. 1952. The Mollusca of Krusadai Island
(in the Gulf of Manaar). I. Amphineura and Gastropoda.
Bulletin of the Madras Government Museum, N. S.,
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pls. 1-33.
Shirley, J. 1912. Additions to the marine Mollusca of
Queensland. Proceedings of the Royal Society of
Queensland, vol. 23, pp. 93-102.
Shopland, E. R. 1896. List of shells collected at Aden in
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Smith, E. A. 1897. Notes on some type-specimens in the
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Solem, A. 1959. Marine Mollusca of the New Hebrides.
Pacific Science, vol. 13, pp. 253-268.
Steele, P. H. 1957. Easter Island Shells. The Nautilus,
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Stockley, G. M. 1928. Report on the geology of the Zan-
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Strausz, L. 1966. Die Miozan-Mediterranen
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Abt. 1, Heft 5, pp. 416-464, pls. 1-6.
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fauna des Roten Meeres und des Golfes von Aden.
Nachrichtsblatt der deutschen Malakozoologischen
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129-206, pls. 13-17, text figs.
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Wilson, B. R. and K. Gillett. 1971. Australian Shells. A. H.
& A. W. Reed, Sydney, 168 pp., 106 pls., text figs.
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Wu, S.-K. 1965b. Comparative functional studies of the
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pp. 211-233, pls. 1-5.
Gastro-
[14-811]
Shout tie ail on
Walter O. Cernohorsky has been a malacolo-
gist and the Curator of Molluscs at the Auckland
Institute and Museum, Auckland, New Zealand,
since 1969. Born in Czecholovakia on June 30,
1927, he emigrated to Fiji in the 1950’s where he
was Chief Surveyor for the Emperor Gold Mining
Company and served as Honorary Conchologist
at the Fiji Museum. In 1968, he received a senior
post-doctoral Research Associateship with the
Smithsonian Institution, Washington, D. C. He is
author of numerous scientific papers on the
marine mollusks of the Indo-Pacific, especially
those of the families Mitridae, Muricidae and
Strombidae. His contributions have appeared in
The Nautilus, The Veliger, the Bulletin of the
Auckland Institute and Museum, and the Revue
Suisse de Zoologie. In 1967 and 1972 he authored
two volumes for amateurs entitled, Marine Shells
of the Pacific. Mr. Cernohorsky is an accom-
plished photographer, an active field collector,
is married, and has two children.
2 Emerson and Cernohorsky
Muricidae
buh
Dr. William K. Emerson is Curator of Mollusks
and Chairman of the Department of Living In-
vertebrates at the American Museum of Natural
History, New York City. His research interests
concern taxonomy and zoogeography of late Cen-
ozoic marine mollusks, especially gastropods and
scaphopods. He has participated in several ex-
peditions to west Mexico and the Caribbean
region. A native of California, Dr. Emerson was
born in San Diego on May 1, 1925, and earned
degrees in zoology from the California State
University at San Diego (A.B. 1948) and the
University of Southern California, Los Angeles
(M.S. 1950). From 1951 to 1955, he served as
Museum Paleontologist at the University of Cali-
fornia, Berkeley, where he received a Ph.D. in
invertebrate paleontology in 1956. He joined the
staff of the American Museum in 1955. Dr.
Emerson is the author of numerous scientific
articles and is co-author of three books designed
to assist amateurs. He is a past president of the
American Malacological Union and of the Wes-
tern Society of Malacologists.
[14-812]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13
Drupa 13
Family Muricidae Rafinesque, 1815
Subfamily Thaidinae Suter, 1909
Key To Subgeneric Groups of Indo-Pacific Drupa
A. Columella singularly folded axially
B. Labial teeth compound,
constricting the aperture
Gash duike vats ankle ena tee Drupa s.str. p. 14-819
BB. Labial teeth singularly arranged or
occasionally in close association, not compound,
aperture not constricted
Prt arenes rete eee ae ar Ricinella p. 14-841
AA. Columella doubly folded axially; margin of outer lip
with 2 digitate processes, aperture narrow,
NOt CoONStricted ........ cee eeeceeeeeeceeeee ees
Subfamily Thaidinae Suter, 1909
The subfamily name Thaidinae has been
granted precedence over the long established
family-group name Purpuridae Menke, 1828,
in Opinion 886 of the International Commis-
sion on Zoological Nomenclature. Although
the Commission has dated Thaidinae Suter
from 1913, Suter erected the family-group
name in 2 prior publications in 1909 (see syn-
onymy).
Authors generally divide the family Muri-
cidae into 7. subfamilies: the Muricinae,
Ocenebrinae, Aspellinae, Thaidinae, Trophon-
inae, Typhinae and Rapaninae. Some authors,
however, consider the Thaidinae worthy of
family rank, but since no scientific evidence
as to important anatomical, morphological or
ecological differences between the Thaidinae
and other closely related muricid groups has
been presented, an elevation of Thaidinae to
family rank appears unwarranted. In such a
closely related species-group as the Muricidae,
with numerous existing species of ‘interme-
diate generic characters, a taxonomic consoli-
dation rather than further subdivision, is indi-
cated.
err Srey oer Te Drupina p. 14-859
Apart from the nominate type-genus Thais
Réding, 1798, the subfamily Thaidinae_ con-
tains such genera as Mancinella Link, 1807,
Acanthina Fischer von Waldheim, 1807, Cymia
Morch, 1860, Purpura Bruguiére, 1789, Nucella
Réding, 1798, Nassa Réding, 1798, Vexilla
Swainson, 1840, Pinaxia H. & A. Adams, 1853,
Xanthochorus Fischer, 1888, Morula Schu-
macher, 1817, Azumamorula Emerson, 1968,
Drupella Thiele, 1925, Neothais Iredale, 1912,
Lepsiella Iredale, 1912, Agnewia Tenison-
Woods, 1878, Cronia H. & A. Adams, 1853, and
other subgeneric groups and fossil genera.
Synonymy—
1828 Purpuracea Menke, Synopsis methodica Mollus-
corum, p. 34 (suppressed by the ICZN in Opinion
886, 1969, Bull. zool. Nomenclature, vol.’ 26, pp.
128-132).
1839 Purpuridae Broderip, Penny Cyclop., vol. 14, p. 321
(suppressed in Opinion 886 of ICZN).
1840 Purpurinae Swainson, Treatise on Malacology, p.
71 (suppressed in Opinion 886 of ICZN).
1909 Thaisidae Suter, Records Canterbury Museum, vol.
1, p. 11; 1909 Suter, Subantarctic Islands of New
Zealand, art. 1, p. 27.
1938 Drupinae Wenz, Handbuch der Palaozoologie, vol
6, pt. 1, pp. 42, 47; 1941 Wenz, ibid., pt. 9, p. 1112.
[14-817]
14 Emerson and Cernohorsky
Subfamily Thaidinae Suter, 1909
Genus Drupa Roding, 1798
Type: Drupa morum Réding, 1798
On shell characters, the group generally
considered as belonging to Drupa s.str. can be
divided into two distinct groups. In the group
of the type-species Drupa morum_ Réding,
which also includes the subspecies D. morum
iodostoma (Lesson), D. ricinus (Linnaeus) and
D. elegans (Broderip & Sowerby), the majority
of the denticles on the outer lip are arranged
as compound, i.e. bifid or trifid teeth which
constrict the aperture posteriorly. In the group
comprising D. rubusidaeus Réding, D. speci-
osa (Dunker) and D. clathrata (Lamarck), the
denticles of the outer lip are singularly situ-
ated as non-compound teeth, which results in
an appreciably wider aperture. For this group
of species the subgeneric name Ricinella
Schumacher, 1817, is available. Although the
teeth on the outer lip of Drupa (Drupina)
grossularia Réding, the type-species of the
subgenus Drupina Dall, are also singularly
arranged, the shell of the species develops
marginal lobate processes and the radula_ has
a greatly modified rachidian radular tooth and
very slender and small lateral teeth.
Drupa differs from Morula in having a more
sub-ovate form, lower spire, longer aperture
and a_ structurally different central radular
tooth. Members of Drupa show a close radular
relationship with most of the thaidine groups.
Muricidae
Synonymy—
1798 Drupa Réding, Museum Boltenianum, p. 55. Type-
species by subsequent designation, Rovereto, 1899:
Drupa morum Réding, 1798.
1807 Canrena Link, Beschr. Nat.-Samml. Univ. Rostock,
p. 126. Type-species by monotypy: Canrena_neri-
toidea Link, 1807 [= Drupa morum Roding, 1798]
(as restricted by Iredale, 1937, and lectotype. des-
ignation by Cernohorsky, 1969).
1810 Sistrum Montfort, Conchyliologie systématique, vol.
2, p. 595. Type-species by original designation:
Sistrum album Montfort, 1810 [= Drupa_ ricinus
(Linnaeus, 1758) ].
1816 Ricinula’ Lamarck, Tableau Encyclopédique — et
Methodique, p. 1. Type-species by subsequent
designation, Children, 1823: Ricinula horida (=
R.horrida Lamarck)
1798].
1822 “Ricinella Lam.”, Bowdich, Elements of Conchology,
vol. 1, p. 40. [? error for Ricinula Lamarck, 1816]
(sole species listed and figured Ricinula horrida
Lamarck, 1816) [non Ricinella | Schumacher,
1816].
1852 Pentadactylus Mérch, Catal. Conchyl. Yoldi, vol. 1,
p. 87. Type-species by subsequent designation,
Baker, 1895: Pentadactylus ricinus Lamarck = Drupa
ricinus (Linnaeus, 1758). [non — Pentadactylus
Schultze, 1760, in Echinodermata; nec Gray, 1845,
in Reptilia].
1855 Ricimula Gould, U.S. Astronom. Exp. South. Hemi-
sphere, vol. 2, p. 263 (error for Ricinula Lamarck,
1516).
1859 Ricinulus Demarest in Chenu, Encycl. Hist. Nat.
Crust. Moll. Zooph., p. 174 (invalid emendation
for Ricinula Lamarck, 1816).
As pointed out by the junior author (Cerno-
horsky, 1969), Suter (1913) is credited by
Dodge (1957) and other authors with the type
designation of Drupa, but Rovereto’s (1899)
designation is earlier.
Although members of _ this
[= Drupa morum Rodding,
genus were
placed until the turn of the present century in
Ricinula,
Lamarck’s there were available
Plate 9. Type-species of subgenera of Drupa Réding. _
Fig. 1. Drupa (Drupa) morum Réding. Namoui reef, Niue
Id. (WOC coll.; 29.8 x 27.4 mm).
Fig. 2. Drupa (Ricinella) rubusidaeus Riding. Suva reef,
Fiji Ids. (WOC coll.; 40.5 x 36.5 mm).
Fig. 3. Drupa (Drupina) grossularia Réding. Pango Point,
Efate Id., New Hebrides (WOC coll.; 28.3 x 28.6 mm).
[14-818]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13
Drupa 15
three valid prior generic names, of which
Drupa Roding, 1798, is the oldest. Link, 1807,
established Canrena as a monotypic genus
with Canrena neritoidea Link, the sole species.
His citation to Martini’s figures include the
species Drupa ricinus (Linnaeus), D. morum
Réding, and D. grossularia Réding. Canrena
neritoidea was restricted by Iredale (1937) to
Drupa morum Roding, as depicted by Mar-
tinis figures 972, 973, on plate 101. The same
figures were designated as the lectotype of
the species Canrena neritoidea Link, by Cern-
ohorsky (1969).
The monotypic genus Sistrum Montfort,
1810, has been erroneously used by many
authors for the species rightly contained in
Morula Schumacher, 1817, which has Drupa
uva Réding, as the type-species. Sistrum al-
bum Montfort, the type-species of Sistrum by
original designation, is a synonym of Drupa
ricinus (Linnaeus). In his discussion of Sistrum
album, Montfort considered the species to
have a white, violet or yellowish aperture, a
misconception common with writers of the
day. Although the violet aperture would indi-
cate the species Drupa morum, Montfort’s fig-
ure of Sistrum album, the specific name itself
and its French, Dutch and Flemish equivalents
of “Le Sistre blanc,” “La mure blanche,”
“wite moerbesie” and “witte moerbeyer,” all
refer to the white-mouthed form of Drupa
ricinus (Linnaeus).
Subgenus Drupa sensu stricto
Shell small to medium, strong, heavy, sub-
ovate; whorls low or conical, often with si-
phonous tubercles, 2 tubercles rarely extended
from the margin as lobate processes; surface
generally sculptured with imbricated — scales;
aperture noticeably constricted posteriorly by
labial teeth; teeth arranged in compound
groups; columella typically with 1 prominent
axial fold, rarely with 2 folds, and with 3-5
prominent oblique plications; radular formula
1-1-1, lateral teeth fang-like, similar in shape,
central tooth typically composed of a large
medial cusp, flanked by a pair of slightly
smaller, bifid to quadrifid cusps, and small,
slender lateral denticles; operculum sub-lunar
to sub-linear, nucleus marginal, marginal cal-
lus well developed, muscle scars few in num-
ber, arranged as gyratory lines.
Members of this genus are typical coral and
reef forms, being confined to the tropical
waters of the Indo-Pacific region, although
some species occur at Easter, Clipperton and
the Galapagos Islands, in the tropical eastern
Pacific.
Drupa morum morum Roding, 1798
(PI. 2, figs. 1-3; Pls. 10, 11)
Range—Red Sea and East Africa to the East-
ern Pacific including Easter and Clipperton Is-
lands (except the Marquesas Islands).
Remarks—The_ purple-colored aperture and
the shorter tubercles distinguish this species
from the white- or yellow-mouthed form of
D. ricinus. The possession of tubercles, the
deeper purple color of the aperture and the
lack of dorsal bands offset this from the Mar-
quesan subspecies D. morum iodostoma (Les-
son, 1840).
Habitat—Intertidal on reef-flats, among rocks
and in crevices; frequently found near the
reef-edge on reefs exposed to strong surf.
Description—Shell 18 to 49 mm (4% to 2
inches) in length, ovately globose, spire acu-
minate, generally inconspicuous’ in adult
stage. Whorls nodose with four rows of short,
strong tubercles. Interstices between tubercles
striated with scalelike striae. Aperture — nar-
row, linear, extending nearly three-quarter the
length of the shell. Outer lip crenulated be-
tween tubercles, dentate on the interior mar-
gin with plaitlike teeth; teeth arranged in an
upper group of four denticles and in a lower
group of three denticles. Two conspicuous
plaits situated immediately above the deep
canal. Inner lip enameled with a callus con-
taining 3 or 4 plaitlike ridges projecting into
the aperture above the canal. Posterior si-
phonal canal elongate, obliquely recurved to-
ward the apex. Color white or grayish, tuber-
cles dark-brown; interior of aperture deep
purple in adults, light purple in immature
specimens. Operculum typical of the group.
Naa
O..mm
Plate 10. Radula of Drupa (Drupa) morum morum Roding.
Half a transverse row; Fiji Ids.
[14-819]
16 Drupa
The central cusp of the rachidian of the
radula is slightly longer than the flanking and
bifid side-cusps; the side-cusps are followed
by 3 to 5 small lateral denticles, exclusive of
the slightly stronger end-cusps. In some speci-
mens, the lateral denticles descend onto the
side-cusps.
Measurements (mm)—(including
specimens with a mature lip).
spines; all
length width
49.1 45.9 S. E. Zanzibar
40.0 38.2 Lectotype of horrida Lamarck
32.4 29.6 Tosa, Japan
27.0 23.7 Viti Levu, Fiji
18.1 17.1 Bikini Atoll, Marshalls
Synonymy—
1684 Buonanni, Rec. ment. oculi anim. test., (3),
fig. 173 only (very poor).
1685 Lister, Hist. Syn. Meth. Conchyliorum, pl. 804,
fig. 13.
1753 Klein, Tent. meth. ostr. nat. cochl., pl. 1, fig. 30.
1758 Nerita nodosa Linnaeus (pars), Systema Naturae,
ed. 10, p. 777 (refers to Lister, Klein and Buon-
anni, (fig. 173) only).
1767 Murex neritoideus Linnaeus (pars), Systema Na-
turae, ed. 12, p. 1219 (refers to Lister, Klein and
Buonanni (fig. 173) only; 1791 Gmelin (pars), Sys-
tema Naturae, ed. 13, p. 3537 (refers to Lister (fig.
13 only), Buonanni, Klein, Knorr and Martini (figs.
972, 973 only) and Seba (fig. 41 only); 1825 Wood,
Index Testaceologicus, p. 123, pl. 26, fig. 47a.
Knorr, Verg. Augen u.Gemiuths, pt. 1, pl. 25,
figs. 5, 6 (good).
1777 “Murex Morum_ globosum” Martini, Syst. Conch-
ylien-Cabinet, vol. 3, p. 280, pl. 101, figs. 972, 973
(East Indies and coast of Coromandel) [non_ bi-
nomial].
1798 Drupa morum Roding, Museum Boltenianum, p. 55
refers to Martini and Knorr) [no locality given];
1913 Hedley, Nautilus, vol. 27, no. 7, p. 80; 1936
Hirase, Coll. Jap. shells, p. 79, pl. 110, fig. 8;
1957 Kaicher, Indo-Pacific Sea Shells, pl. 4, fig. 3;
1960 Hertlein, Veliger, vol. 3, no. 1, p. 8 (Gala-
pagos and Clipperton Ids.); 1961 Rippingale & Mc-
Michael, Queensland and Gt. Barrier reef shells,
pl. 13, fig. 5; 1965 Arakawa, Venus: Jap. Journ.
Malacology, vol. 24, no. 2, p. 114, pl. 13, figs. 3, 4
(radula); 1965 Wu, Bull. Inst. Zool. Acad. Sinica,
vol. 4, p. 98, text fig. 19 (radula); 1967 Orr Maes,
Proc. Acad. Nat. Sci. Philadelphia, vol. 119, no. 4,
p. 129; 1969 Cernohorsky, Veliger, vol. 11, no. 4,
p. 298, pl. 47, fig. 7 (shell), text fig. 4 (radula); 1970
Salvat, Cahiers du Pacific, no. 14, p. 46; 1971 Wilson
& Gillett, Australian Shells, p. 92, pl. 61, fig. 2; 1971
Kay, Pacific Science, vol. 25, pp. 266, 275.
1807 Canrena_ neritoidea Link, Beschr. Nat.-Samml. Univ.
Rostock, 3 Abth., p. 126 (refers to Martini, pl. 101,
figs. 972, 973—designated as lectotype figures by
Cernohorsky, 1969) [no locality given].
1816 Ricinula horrida Lamarck, Tabl. Encycl. Methodique,
p. 1, pl. 395, figs. la, b (no locality given); 1822
Lamarck, Hist. nat. anim. s. vertebres, vol. 7, p.
231 (Indian Ocean); 1823 Sowerby, Genera Rec.
foss. shells, pt. 18, pl. 235, fig. 1; 1842 Reeve,
Conchologia Systematica, vol. 2, p. 215, pl. 156,
fig. 1; 1846 Reeve, Conchologia Iconica, vol. 3,
pl. 1, fig. 3; 1859 Chenu, Manuel Conchyliologie,
1768
Emerson and Cernohorsky
Muricidae
vol. 1, p. 168, fig. 814; 1880 Tryon, Manual Con-
chology, vol. 2, p. 184, pl. 56, figs. 201, 202; 1933
Dautzenberg & Bouge, Journal de Conchyliologie,
vol. 77, p. 238.
1817 Ricinella violacea Schumacher, Essai Nouv. Sys-
téme, p. 240 (refers to Martini, op. cit.) [no local-
ity given].
1823 Ricinula horida (sic) Children, Quart. Journ. Sci.
Lit. & Arts, vol. 16, p. 56, pl. 5, fig. 189.
1832 Purpura horrida Lamarck, Blainville, Nouv. Ann.
Mus. d’Hist. Nat. Paris, ser. 3, vol. 1, p. 208; 1833
Quoy & Gaimard, Voyage L’Astrolabe, vol. 2,
p. 576, pl. 39, figs. 1-3 (animal and operculum);
1835 Kiener, Spéc. gen. icon. coq. viv., vol. 8, p. 8,
pl. 1, fig. 1 (animal).
1850 Sistrum horridum M. Gray, Figs. Moll. Animals,
vol. 4, p. 70, pl. 96, fig. 11; 1911 Schepman, Si-
boga-Expeditie, vol. 49d, p. 355; 1952 Morris, Field
Guide shells Pacific coast and Hawaii, pl. 39,
fig. 5.
1852 Ricinula (Pentadactylus) globosa Mart., Morch, _ Cat.
Conchyl. Yoldi, vol. 1, p. 88 (synonymized with
R. horrida Lamarck and Drupa morum Roding).
1853 Pentadactylus (Pentadactylus) globosus H. & A.
Adams, Genera Rec. Mollusca, vol. 1, pp. 129, 130,
pl. 13, fig. 6 (animal).
1908 Pentadactylus (Pentadactylus) horridus | Lamarck,
Horst & Schepman, Cat. Syst. Moll. Mus. Hist. Nat.
Pays-Bas, vol. 13, p. 157.
1938 Drupa (Drupa) morum Roding, Adam & Leloup,
Mém. Mus. Roy. d’Hist. Nat. Belg., vol. 2, fasc.
19, p. 164.
Types—Since Roding’s types are probably no
longer extant, we designate the specimen de-
picted on plate 101, figs. 972, 973 in Martini,
as the lectotype of Drupa morum Roding (pl.
11 fig. 1). Two syntypes of Ricinula horrida
Lamarck, are in the Muséum d Histoire Natur-
elle, Geneva; the specimen measuring 40.0
mm in length, no. 1101/12/1, is here selected
as the lectotype of R. horrida. The type-speci-
men of Ricinella violacea Schumacher, could
not be located in the University Zoological
Museum, Copenhagen. From the two localities
mentioned by Martini for D. morum s. s. we
select the first-mentioned “East Indies” as the
Plate 11. Drupa (Drupa) morum morum Roding.
Fig. 1. Lectotype figure from Martini, 1777, Syst. Con-
chylien-Cabinet, vol. 3, pl. 101, fig. 972.
Fig. 2. Lectotype of Ricinula horrida Lamarck (MHNG no.
1101/12/1; 40.0 x 38.2 mm).
[14-820]
May 30, 1973
type locality, and restrict it further to Java,
Indonesia (specimens in AMNH and ANSP).
Nomenclature—This species was well known
to early naturalists but was confused by Lin-
naeus with Thais nodosa and T. nodosa ascen-
sionis Quoy & Gaimard (= meretricula Roding).
In his 12th edition of the “Systema Naturae,”
Linnaeus re-named the species Murex neri-
toideus, and cited the same erroneous figures
which depict Drupa morum. However, in Lin-
naeus’ personal copy of the 12th edition of the
“Systema Naturae,” the complete synonymy
of the 10th edition description of Nerita nodosa
has been cancelled and the following words were
added: “Labium interius punitis 2 maculatum.”
Records—RED SEA: Lahonel Beach, Gulf of Aqaba,
Israel (Lamy, 1938, p. 56); Berenice, Egypt (ANSP); Mah-
mud reef, Jiddah, Saudi Arabia (AMNH). EAST AFRICA:
SE Point, Isla di Serpenti, Chisimaio, Somalia (AMNH);
Pangavini Id., 10 mi. NNE_ Dar-es-Salaam, Tanzania
(MCZ); Diani Beach, Kenya (Heinicke, 1970, p. 7); Moz-
ambique (USNM). ZANZIBAR: Chumbe Id.; Kiwengwa;
Ras Nungwe (all ANSP). SEYCHELLES: (AMNH); Cous-
in Id. (ANSP). MADAGASCAR: S. side of Nossi Iranja,
32 mi. SW of Nossi Bé; Flacourt, Fort Dauphin; 2 mi. NE
Point Ansiraikiraiky (all MCZ). MAURITIUS: 1 mi. ESE
of Souillac; Pointe Fayette (both ANSP); near Port Louis
(MCZ); Point Pimente, N. side Arsenal Bay (Powell coll.).
REUNION ISLAND: (ANSP). CHAGOS ARCHIPELAGO:
(Melvill, 1909, p. 104). MALDIVE ISLANDS: Imma Id., SE
North Male Atoll; Fadiffolu Atoll; Tiladummati Atoll (all
ANSP), LACCADIVE ISLANDS: (Hornell, 1922, p. 217).
INDIA: Pamban and Shingle Id., Gulf of Manaar (Satya-
murti, 1952, p. 160). CEYLON: 12 mi. N. of Trincomalee
(AMNH); Hikkaduwa (ANSP). THAILAND: Loam Seng,
S. of Laam Son, Phuket Id. (ANSP). VIETNAM: Cam Ranh
Bay (ANSP). COCOS-KEELING ISLANDS: N. end of Pulo
Siput; West Id., Cocos Id. (both ANSP). CHRISTMAS
ISLAND: (Tomlin, 1935, p. 79). INDONESIA: Jesselton
district, N. Borneo; Pulan Boenta, off Acheh Head, N. W.
Sumatra; Sulau Bay, Batu group, off Sumatra; Mega,
Mentawai Ids., S. :
W.Sumatra (all USNM); Queen’s Bay,
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13
Drupa 17
off Sukabumi, Java (AMNH); Keledjitan, Bantam, Java
(ANSP); Batjan Id.; Manipa Id., W. of Halmahera Id._;
Mandidi Id.; Ambayana Id., all Moluccas (all MCZ).
PHILIPPINES: Corregidor Id.; Manila Bay, Luzon Id.;
Calapan, Mindoro Id.; Gigmoto, Catanduanes Id.; Cuyo
Id., Palawan group (all ANSP); Panay; Silino Id., Minda-
nao Id. (both USNM); Sanga Sanga Id., Sulu Archipelago
(ANSP). FORMOSA (Taiwan): (AMNH); Suo; Karenko;
Botan-wan; Ryukyu-Syo; Hoko (all Kuroda, 1941, p. 111).
RYUKYU ISLANDS: Hyakuna reef (AMNH); Yomitan reef,
Okinawa: Bolo Point; Kuzu-Saki, all Okinawa (all ANSP);
NE coast of Iheya Shima (ANSP); Miyako (FMNH). JA-
PAN: Kikaiga Shima, Osumi; Oshima Osumi (MCZ;
USNM); Tosa (ANSP); Kii, Honshu (FMNH). MARIANAS:
Maung: Agrihan (both USNM); Saipan; Tinian; Guam (all
AMNH). PALAU ISLANDS: Babelthuap (ANSP); Koror;
Eil Malk (both USNM). CAROLINE ISLANDS: Kayangel;
Angulpelu; Gorokoru; Ngargersiul (all ANSP); Yap
(USNM); UWlithi; Ifalik; Elato; Satawal; Ponape; Kaping-
amarangi (all ANSP). MARSHALL ISLANDS; Eniwetok
Atoll; Rongelap Atoll; Majuro Atoll; Ujae Atoll; Uterik
Atoll (all USNM):; Bikini Atoll; Arno Atoll (both AMNEFHI).
WAKE ISLAND: (AMNH; ANSP; MCZ). NEW GUINEA:
Schouten Ids.; Wewak:; Makiri reef, Unea Id. (all ANSP);
2 mi N. of Gusika, 13 mi. N. of Finschhafen (MCZ). AD-
MIRALTY ISLANDS: Koruniat Id. (ANSP). AUSTRALIA:
Queensland: Herald Cay, Coral Sea; Pipon reef near Cape
Melville (both AMS): Wilson Id., Capricorn group (MCZ);
Lady Elliot Id. (ANSP; AIM); Heron Id. (AIM); Tryon Id.,
Capricorn group (Powell coll.); | Northern Territory: Dar-
win (FMNII); New South Wales: Middleton reef, off coast
of New South Wales (Iredale, 1937, p. 256); Lord Howe
Id. (AMS): West Australia: Barrow Id. (Wilson & Gillett,
1971, p. 92). NEW BRITAIN: Rabaul Harbour (ANSP).
NEW IRELAND: Kavieng (AMNH). SOLOMON ISLANDS:
NW. Bougainville Id. (ANSP); Lutee, Choiseul Id.; Ataa,
N. Malaita) Id.; Lunga, Guadalcanal; Bellona Id. (all
AMNIIL); Vikopia Id. (AIM); Bougainville; Bumana (both
Powell coll). NEW) HEBRIDES: Tongoa, Espiritu) Santo
Id. (MCZ); Bushmen’s Bay, E. Malekula Id. (AIM); Tanna
Id. (DM). NEW CALEDONIA: Touho (AMNII); Isle of
Pines (MCZ). LOYALTY ISLANDS: Lifu) (USNM). FIJI
ISLANDS: Savusavu, Vanua Levu (AMNH); Korolevu,
S. Viti Levu (ANSP); Nananu-i-Ra Id., N. Viti Levu (WOC
coll.); Ongea Levu, Lau Ids. (USNM). GILBERT IS-
LANDS: Onotoa Atoll (MCZ); Kingsmill Id. (USNM);
Nauru Id. (Iedley, 1903, p. 4). ELLICE ISLANDS: Funa-
futi (AMS: AIM). WALLIS ISLANDS: Nukuhifala (USNM).
TONGA | Islands: Hlufagalupe, Tongatapu (ANSP); Ni-
uafo’ou Id. (USNM). SAMOA ISLANDS: Asau_ Harbour,
160° 180°
160°
T
140° \20° 100° 90°
T r T
Plate 12. Geographical distribution of Drupa (Drupa) morum
morum Réding, and its Marquesan subspecies D. (D.)
(Lesson). Open circles are literature
morum iodostoma
records.
(14-821]
18 Drupa
Savaii; Pago Pago, Tutuila (both USNM); Apia, Upolu
(AMNH; AIM). NIUE ISLAND: Alofi (USNM); Oneone
reef; Namoui reef (both AMNH). PHOENIX ISLANDS:
Canton Id. (ANSP). COOK ISLANDS: Bird’s Id.; North Id.,
both Palmerston Atoll (both USNM); Aitutaki; Mauke;
Rarotonga; Mangaia (all USNM); Kopuano Passage, Aitu-
taki (AIM). PENRHYN ISLAND: (AMNH; AIM). AUSTRAL
ISLANDS: Tubuai; Raivavae; Rurutu (all USNM). SO-
CIETY ISLANDS: many localities on Bora Bora; Huahine;
Moorea; Tahiti (all USNM); Venus Point, Tahiti (AMNH).
TUAMOTU ISLANDS: Tikahau Atoll; Vahitahi; Fakarava;
Nengonengo; Makatea (all USNM). GAMBIER ISLANDS:
Mangareva (USNM). LINE ISLANDS: Palmyra _ Id.
(USNM):; Fanning Id. (AMS); Caroline Id.; Flint Id. (both
ANSP); Jarvis Id. (Powell coll.); Kingman reef; Washing-
ton Id.; Christmas Id. (all Kay, 1971, p. 275). HAWAIIAN
ISLANDS: Kamilo, Kauai; Hanauma Bay, Oahu; Pukoo,
Molokai; Honokowai reef, Maui; Hilo, Hawaii (all USNM);
Kona, Hawaii (AMNH); Koko Head, Oahu; Waikiki, Hon-
olulu (both Powell coll.). PITCAIRN ISLAND: (AMNH).
EASTER ISLAND: (Steele, 1957, p. 112). CLIPPERTON
ISLAND: (Hertlein & Allison, 1960, p. 15).
Fossil records—HAWAIAN ISLANDS: Pleistocene: Oahu
(Ostergaard, 1928, p. 6; Kosuge, 1969, p. 786, pl. 5, fig.
93).
Drupa morum iodostoma (Lesson, 1840)
(PI. 2, figs. 4, 5; Pl. 13)
Range— Marquesas Islands.
Remarks—The lack of tubercles, the more
quadrate shape, the dark dorsal bands and the
pinkish violet aperture distinguish this uncom-
mon subspecies from the nominate species
Drupa morum Roding.
Early records from both literature and spec-
imens are from numerous localities in the
western Indo-Pacific, but reliable data indi-
cate this taxon to be restricted to the Mar-
quesa Islands. Records based on old _ collec-
tions in the National Museum of Natural His-
tory, Washington, and the Museum of Com-
parative Zoology, Harvard, can be confidently
dismissed as being based on erroneous local-
ity data, probably copies from locality indica-
tions given in old literature; these localities
are “Malacca,” “New Zealand,” “Madagascar”
and “Fiji Islands.” However, there are several
records of “D. iodostoma” from neighbouring
areas to the Marquesas, such as Palmyra Island
in the Line Islands (SDNHM), Venus Point,
Tahiti (AMNH) and Fakarawa Atoll, Tuamo-
tus (ANSP); in addition, the subspecies has
also been reported in literature from Marutea
Island (Couturier, 1907) and Makatea Island
(Boettger, 1918), both in the Tuamotu Archi-
pelago. The literature records are suspected
Emerson and Cernohorsky
Muricidae
misidentifications, while the actual specimens
in the mentioned institutions have either been
obtained through secondhand or have been
documented by persons who also collected in
the Marquesas Islands apart from the Tuamo-
tus. Confirmed, recent collections suggest,
therefore, that D. iodostoma is an allopatric
subspecies of D. morum, and is endemic to
the Marquesa Islands where D. morum is not
known to occur.
Habitat—In surge channels cut into exposed
volcanic shores (G. D. Stout, personal com-
munication), and in rocky crevices at a depth
of 10 feet.
Description—Shell 32 to 47 mm (about 1% to
2 inches) in length, quadrately globose, spire
very short and acuminate. Whorls faintly ax-
ially plicate, spirally ribbed by five wide,
slightly raised ribs which are more __pro-
nounced toward the margin. Interstices be-
tween ribs striated with 3 lirations. Aperture
narrow, linear, extending nearly three-quarter
the length of the shell. Outer lip crenulated
between ribs, inner margin dentate; teeth
compound, 3 to 4 denticles in the upper tooth,
2 to 3 denticles in the lower tooth. Above the
anterior canal are two conspicuous plaits. Col-
umella with three to four heavy plications pro-
jecting into the aperture. Posterior siphonal
canal elongate, obliquely curved toward the
apex. Color cream, spiral ribs brownish black,
intersticial spiral threads reddish brown; aper-
ture pinky-violet.
The radula is similar to Drupa morum; the
rachidian has bifid side-cusps and 4 or 5 lat-
eral denticles. The radula of “Drupa_ iodos-
toma” as figured by Cooke (1919) from the
“Sandwich Islands” [= Hawaiian Ids.], is
probably based on Hawaiian specimens of
D. morum.
Measurements (mm)—(all specimens with a
mature lip).
length width
47.0 43.0 Nukuhiva, Marquesas Ids.
38.0 36.5 Nukuhiva, Marquesas Ids.
36.0 32.7 Tahuata, Marquesas Ids.
30.3 27.0 Tahuata, Marquesas Ids.
Synonymy—
1840 Purpura (Ricinula) iodostoma Lesson, Rev. Zool.
Soc. Cuvierienne, vol. 3, p. 355 (“New Zealand”
= error!).
1842 Purpura iodostoma Lesson, Guerin’s Magasin de
Zoologie, vol. 4, p. 58, pl. 58.
[14-822]
May 30, 1973
LO
Plate 13. Radula of Drupa (Drupa) morum iodostoma (Les-
son). Half a transverse row; Tahuata, Marquesa Ids.
1846 Ricinula iodostoma Lesson, Reeve, Conchologia
Iconica, vol. 3, pl. 1, figs. 4a, b; 1860 Reeve, Ele-
ments of Conchology, vol. 1, p. 82, pl. 7, fig. 32;
1880 Tryon, Manual of Conchology, vol. 2, p. 184,
pl. 56, fig. 199.
1888 Ricinula jodostoma (sic) Lesson, Paetel, Catalog der
Conchyl.-Sammlung, vol. 1, p. 143 (New Zealand
[erroneous] ).
1908 Pentadactylus (Pentadactylus) iodostomus Lesson,
Horst & Schepman, Cat. Syst. Moll. Mus. Hist.
Nat. Pays-Bas, vol. 13, p. 154.
1913 Drupa iodostoma Lesson, Hedley, The Nautilus, vol.
27, no. 7, p. 80; 1957 Kaicher, Indo-Pacific Sea
Shells, pl. 4, fig. 5 (Malaysia and Melanesia [=
error!]); 1965 Hertlein, Occ. Papers California
Academy of Sciences, no. 49, p. 2, figs. 1, 2 (Mar-
quesas Islands).
1918 Ricinula jodostoma_ (sic) Lesson, Boettger, Abh.
Senckenb. Naturfr. Gesellschaft, vol. 36, (3), p.
298 [Probably misidentified Drupa morum Rod-
ing].
Types—The type-specimen of Drupa_ iodos-
toma (Lesson) is probably in the Muséum Na-
tional d’Histoire Naturelle, Paris. The given
locality “New Zealand” is erroneous, as_ the
species does not live there. Since the type-
specimen was collected during the voyage of
the “Vénus,” which visited the Marquesas
Phi
0.1mm
LSA
5.0 mm
Plate 14. Drupa (Drupa) ricinus ricinus (Linnaeus)—white
mouthed form; Fiji Islands.
Fig. A. Radula; half a transverse row.
Fib. B. Part of rachidian of radula showing variation in the
number of accessory denticles on the side-cusps.
Fig. C. Penis.
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 19
after the 14th of August, 1838, and also called
at the Bay of Islands, New Zealand, during
October 1838 (Chamberlin, 1960, p. 67), the
erroneous locality is obviously due to a mix-up
of specimens during the voyage. Since reliable
records are known only from the Marquesas,
we designate Taiohae, Nukuhiva Island, Mar-
quesa Islands, as the type locality of D.
morum iodostoma.
Records—MARQUESAS ISLANDS: Hana Nui and Haavie
Bays, Ua Huka Id.; Eiao Id. (all ANSP); Taiohae, Nuku-
hiva Id. (USNM; AMNH); N. side Hana Moe Noe Bay,
Tahuata (USNM).
Drupa ricinus ricinus (Linnaeus*, 1758)
(Pl. 2, figs. 6-5, 11; Pls. 14, 15, 16)
Range—From East and South Africa to the
Eastern Pacific, at Easter, Clipperton and Gal-
apagos Islands.
Remarks—Differs from D. morum, which has
a proportionally larger shell, in having longer
spines, a smaller size and in lacking the pur-
ple or mauve apertural coloration. D. elegans
(Broderip & Sowerby) differs mainly by _ pos-
sessing a continuous, red-brown line — encir-
cling the aperture; it is restricted to Polynesia
where it is found with the present species.
°It is the editorial policy to allow authors to use their own
preference in spelling this name. Linnaeus or Linne.
0.1mm
B
Plate 15. Drupa (Drupa) ricinus ricinus (Linnaeus)—yellow
mouthed form; Fiji Islands.
Fig. A. Radula; half a transverse row.
Fig. B. Part of rachidian of radula showing variation in the
side-cusps and lateral denticles.
[14-823]
20 Drupa
Commonly, D. ricinus may have a faint orange
or yellow, diffused line encircling the aper-
ture.
Populations with large shells, attaining
nearly 40 mm in length, are present in the Red
Sea. This form appears to be geographically
isolated from the smaller-shelled, nominate
subspecies, and it is recognized herein as a
new subspecies Drupa ricinus hadari.
Habitat—Lives exposed, or in water up to
approximately 10 feet deep, on or under
rocks, sand, weed or coral, on exposed wave
benches or fringing reefs. At Clipperton Island
the species has been collected to a depth of
22 fathoms.
Description—Shell 17 to 35 mm (about % to
1% inches) in length, solid, obovoid, spire
short, reduced, obscure in adult specimens,
body whorl three-quarter the length of the
shell. Whorls ribbed with five rows of sharp
spines; spines variable in development, but
higher near the margin of the aperture. Inter-
stices between spines striated with scale-like
striae. Aperture linear, constricted by the
teeth and callus. Apertural dentition as in
D. morum with features less prominent. Color
white or grayish; spines dark-brown to dark-
gray, especially toward the tips; mouth white
or with a wide, broken, diffused, orange to
yellow line extending around the outer lip and
neighbouring canal area. Operculum typical
for the group.
In the radula, the side-cusps of the rachid-
ian are short or moderately long, bifid to
quadrifid, and are followed by 2-3. side-
denticles, exclusive of the end-cusps.
Measurements (mm)—(including spines; all)
specimens With a mature lip).
length width
32.3 27.8 Kagoshima, Japan
29.5 25.5 Lectotype of arachnoides
Lamarck
25.6 27.0 S. Viti Levu, Fiji Ids.
23.0 25.0 Probable holotype of
ricinus Linnaeus
22.8 21.0 Penrhyn Id.
17.2 16.3 Samarai, Papua
Synonymy—
1705 Rumphius, Amboinsche Rariteitkammer, pl. 24,
fig. E.
1742 d’Argenville, L’Histoire Naturelle ...., pl. 17,
fig. A.
1742 Gualtieri, Ind. test. Conchyliorum, pl. 28, fig. N
(poor).
Emerson and Cernohorsky
Muricidae
1758 Murex ricinus Linnaeus, Systema Naturae, ed. 10,
p. 750 (refers to Rumphius and Gualtieri, op. cit.)
[Asiatic Ocean] (yellow and white forms).
1758 Murex hystrix Linnaeus, Systema Naturae, ed. 10,
p. 750 (refers to d’Argenville, op. cit.) [no locality
given] (juvenile specimen); 1855 Hanley, Ipsa Linn.
Conch., p. 294.
777 “Murex Morum globosum” Martini, Syst. Conchyl-
ien-Cabinet, vol. 3, p. 280, pl. 102, figs. 976, 977
(non binomial) [yellow form].
1798 Drupa tribulus Roding, Museum Boltenianum, p. 55
(refers to Rumphius and Gualtieri, op. cit.) [no
locality given].
1798 Drupa_ rubuscaesius Roding, ibid., p. 55 (refers to
Martini, op. cit.) [no locality given] (yellow form).
1810 Sistrum album Montfort, Conchyliologie Systemat-
ique, vol. 2, p. 595, fig. on p. 594 (white form)
[no locality given].
1816 Ricinula arachnoides Lamarck, Tableau Encyclo-
pedique Méthodique, p. 1, pl. 395, figs. 3a, b (no
locality given) [yellow form]; 1822 Lamarck, Hist.
nat. anim. s. vertebres, vol. 7, p. 232 (Indian
Ocean); 1823 Sowerby, Gen. Rec. fossil shells,
pt. 15, pl. 235, fig. 5 (juvenile specimen); 1842
Reeve, Conchologia Systematica, vol. 2, p. 215,
pl. 256, fig. 5 (juvenile specimen); 1846 Reeve, Con-
chologia Iconica, vol. 3, pl. 1, fig. 5; 1859 Chenu,
Manuel de Conchyliologie, vol. 1, p. 168, fig. 812.
1831 Murex neritoideus Mawe, Wodarch’s Intr. Conchol-
ogy, pl. 3, fig. 43 (yellow form) [non Linnaeus,
1767].
1832 Purpura albo-labris Blainville, Nouv. Ann. d Hist.
Nat. Paris, ser. 3, vol. 1, p. 208, pl. 9, fig. 5 (Trin-
comalee, Ceylon) [white form]; 1835 Kiener, Spéc.
gen. icon. coq. viv., vol. 8, p. 12, pl. 1, fig. 2.
1832 Purpura arachnoides Blainville, ibid., p. 209 (yel-
low form); 1833 Quoy & Gaimard, Voyage L’Astro-
labe, vol. 2, p. 579, pl. 39, figs. 17-19 (animal and
operculum); 1835 Kiener, Spéc. gén. icon. coq. viv.,
vol. 8, p. 10, pl. 1, figs. 3, 3a; 1848 Krauss, Stida-
frik. Mollusken, p. 115 (Natal).
1850 Sistrum arachnoides Lamarck, M. Gray, Figs. Moll.
Animals, vol. 4, p. 70, pl. 96, fig. 2.
1853 Pentadactylus ricinus Linnaeus, H. & A. Adams, Gen.
Rec. Mollusca, vol. 1, p. 130; 1875 Troschel, Gebiss
der Schnecken, vol. 2, p. 134, pl. 13, fig. 5 (radula).
1859 Ricinula albolabris Blainville, Chenu, Manuel de
Conchyliologie, vol. 1, p. 168, fig. 812.
1880 Ricinula ricinus Linnaeus, Tryon, Manual Conchol-
ogy, vol. 2, p. 184, pl. 56, fig. 200 and pl. 57, figs.
204, 206, 212; 1933 Dautzenberg & Bouge, Journal
de Conchyliologie, vol. 77, p. 240.
1884 Pentadactylus arachnoides Lamarck, Fischer, Man-
uel de Conchyliologie, fasc. 7, p. 646, pl. 6, fig. 9.
1911 Sistrum ricinus Linné, Schepman, Siboga-Expeditie,
vol. 49d, p. 354.
1913 Drupa ricinus Linne, Hedley, Nautilus, vol. 27, no. 7,
p. 80; 1937 Hertlein, Proc. Americ. Phil. Society,
vol. 78, no. 2, p. 308, pl. 1, figs. 5, 6 (Clipperton
and Galapagos Ids.); 1967 Orr Maes, Proc. Acad.
Nat. Sci. Philadelphia, vol. 119, no. 4, p. 129; 1969
Cernohorsky, Veliger, vol. 11, no. 4, p. 299, pl. 47,
figs. 8, 8a (shell), text figs. 5, 6 (radula); 1970
Salvat, Cahiers du Pacifique, no. 14, p. 46; 1971
Kay, Pacific Science, vol. 25, pp. 266, 275; 1971
Wilson & Gillett, Australian Shells, p. 92, pl. 61,
figs. 3, 3a.
1915 Drupa rubus-cestus Dall, Smithsonian Inst. Publ. no.
2360, p. 29 (refers to Réding, 1798, p. 55, species
695 = D. tribulus Roding).
1929 Drupa (Drupa) ricinus Linnaeus, Thiele, Handb.
syst. Weichtierkunde, vol. 1, p. 295; 1938 Adam &
[14-824]
May 30, 1972
Leloup, Mém. Mus. Roy. d’Hist. Nat. Belg., vol. 2,
fasc. 19, p. 164.
1933 Ricinula ricinus var. arachnoides Lamarck, Dautzen-
berg & Bouge, Journal de Conchyliologie, vol. 77,
p. 240.
1960 Drupa ricina Linnaeus, Hertlein, Veliger, vol. 3, no.
1, p. 8; 1965 Arakawa, Venus: Jap. Journ. Malacol-
ogy, vol. 24, no. 2, p. 115, pl. 13, fig. 1 (radula);
1965 Wu, Bull. Inst. Zool. Acad. Sinica, vol. 4,
p. 98, text fig. 18 (radula); 1965 Wu, Malacologia,
vol. 3, no. 2, p. 211, text figs. (anatomy).
1960 Drupa_ ricina forma albolabris Blainville, Hertlein,
Veliger, vol. 3, no. 1, p. 8.
1965 Drupa albolabris (Blainville), Arakawa, Venus: Jap.
Journ. Malacology, vol. 24, no. 2, p. 114, pl. 13,
fig. 2 (radula).
1965 Drupa arachnoides Lamarck), Wu, Bull. Inst. Zool.
Acad. Sinica, vol. 4, p. 98, text fig. 18 (radula).
Types—The probable — type-specimen _ of
Drupa ricinus (Linnaeus) [Pl. 16, fig. 2], is in
the Linnean collection of the Linnean Society,
London. This particular specimen is the yel-
low-spotted form, which has a no. 540 written
on the columella (the number of Murex ricinus
in the 12th edition of the “Systema Naturae’”).
Two additional, undocumented specimens are
also in the collection, an adult of the white
form and a juvenile specimen. Five syntypes
of Ricinula arachnoides Lamarck, are in the
Muséum d'Histoire Naturelle, Geneva, and the
29.5mm long syntype, no. 1101/15/4, is here
selected as the lectotype (PI. 16 fig. 4). The
type-specimen of Purpura albolabris Blain-
ville, is probably in the Muséum _ National
@Histoire Naturelle, Paris. The type locality
of D. ricinus is “Asiatic Ocean,” which is here
restricted to Ceylon (specimens in MCZ, ANSP
and AMNH).
Nomenclature—Much_ confusion has existed
regarding the identity of this taxon. Lamarck,
1822, considered Murex hystrix Linnaeus, to be
the pink-apertured species which is correctly
known as Drupa_ rubusidaeus Roding. This
misconception was continued by many early
authors. Actually, as is pointed out by Hanley
(1855), Linnaeus referred Regenfuss’ (1758)
excellent colored figure of Drupa rubusidaeus
Réding to Murex hippocastanum in his 12th
edition of the “Systema Naturae,” and not to
Murex hystrix. In the 10th edition, the short
diagnosis of Murex hystrix was accompanied
by only one figure reference, pl. 17, fig. A of
d’Argenville, 1742. This engraving (see Pl. 16
fig. 1) unmistakeably represents the dorsal as-
pect of Drupa ricinus as indicated by the long,
obliquely slanted spines which characterize
some forms of this species. In the description
Linnaeus states: “apertura edentula_ repanda,”
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa All
Plate 16. Drupa (Drupa) ricinus ricinus (Linnaeus).
Fig. 1. Lectotype figure of Murex hystrix Linnaeus, from
d’Argenville, 1742, L’Histoire Naturelle . Conchyli-
ologie, pl. 17, fig. A.
Fig. 2. Probable type-specimen of Murex ricinus Linnaeus,
from the “Asiatic Ocean” (Linnean Society of London
coll.; 23.0 x 25.0 mm).
Fig. 3. Lectotype figure of Drupa rubuscaesius Réding,
from Martini, 1777, Syst. Conchylien-Cabinet, vol. 3,
pl. 102, tig. 976.
Fig. 4. Lectotype of Ricinula arachnoides Lamarck (MHNG
no. 1101/15/4; 25.5 x 29.5 mm).
an indication that this species was based on
specimens in which the dentate lip has not yet
developed. Thus it can be safely assumed that
Murex hystrix Linnaeus, is nothing more than
a juvenile of Drupa ricinus Linnaeus.
At the present time, some students recognize
two species, the form with a white aperture as
Drupa albolabris (Blainville) and the yellow-
orange spotted form as D. arachnoides (Lam-
arck). The latter should not be confused with the
yellow-mouthed D. grossularia Réding. In a de-
tailed sampling of D. ricinus in the Fiji Islands
by the junior author (Cernohorsky, 1969), both
[14-825]
22 Drupa
forms were found not only to be sympatric in var-
ious localities, but were sharing the same rock in
many instances. No sexual dimorphism was ob-
served in either color form, and no differences in
either living animal, radula, color of the stomach
pouch or the penis were apparent in either form.
The number of accessory side-denticles in the
rachidian of the radula varied from 3 to 5 in both
color forms. Both color forms of D. ricinus have
been collected together at various other Indo-
Pacific localities. Hertlein and Allison (1960) re-
corded the presence of both color forms of
D. ricinus at Clipperton Island, an isolated coral
atoll in the eastern Pacific, located about 670
miles southwest of Acapulco, Mexico. The white-
apertured form was reported to be living among
boulders and coral debris on the outer parts of
the reef flats, and off the edge of the reef flats,
in coral and coral rubble, to a depth of at least
130 feet, the lower limit of collecting by their
SCUBA divers. Although the white-apertured
form was found in abundance, only abraded
specimens of the yellow-orange form were on the
beach and off shore at a depth of 70 feet. In the
New Hebrides the white-mouthed form only was
found by the junior author.
Records—(inserted in brackets: y = yellow-spotted form,
w = white form). GULF OF ADEN: Aden (Shopland, 1896,
p. 220). EAST AFRICA: Isla di Serpenti, Chisimaio, Somalia
(AMNH; yw); Port Amelia, Mozambique (AMNH; y); Kend-
wa Id., 4 mi. ESE Dar-es-Salaam, Tanzania; Pangavini Id.,
10 mi. NNE Dar-es-Salaam, Tanzania; Ras Kankadya, 6 mi.
N. Dar-es-Salaam, Tanzania (all MCZ; y); Diani Beach,
Kenya (AMNH; USNM; y); Mombasa, Kenya (AMNH; y).
ZANZIBAR: (AMNH; w); Ras Nyngwe; Chumbe Id.; Jembi-
ani, 5 mi. S. Paje; Kiwenga (all ANSP; y). SOUTH AFRICA:
Umtwalnmi, 22 mi. N. Port Shepstone, Natal; East London
(both ANSP; y). SEYCHELLES: Loraie Bay, Curiense Id.;
Anse aux Pins (both ANSP; y); Coetivy Id.; Praslin Id. (Mel-
zo” 40° 60° 80° to0° zoe t40e
Emerson and Cernohorsky
Muricidae
vill, 1909, p. 103). MADAGASCAR: Ambodifototra, Isle St.
Marie (MCZ; w); N. E. of Pointe Antsiraikiraiky, NW Isle
St. Marie; Ambariobe, S.E. Nossi Bé; Grande Recife, Tulear;
Pointe Ibanona, Fort Dauphin (all MCZ; y). REUNION:
(ANSP; w). MAURITIUS: Pointe Fayette; S.W. Port Louis
(both ANSP; w); 1 mi. NW Black River (ANSP; y). MAL-
DIVE ISLANDS: Ongu Id., N. Malosmadulu Atoll; Wala Id.,
Nilandu Atoll; Fadifolu Atoll (all ANSP; y); Imma Id., N.
Male Atoll (ANSP; w). CEYLON: (MCZ; w); Pointe de Gal-
lett (ANSP; y); 12 mi. N. of Trincomalee (AMNH,; y). ANDA-
MAN ISLANDS: (Melvill & Sykes, 1899, p. 222) THAI-
LAND: Laam Seng, 1 mi. S. Laam Son, Phuket Id. (ANSP;
y); Goh Huyong, Similan Ids. (USNM; y). VIETNAM: Con
Son Ids. (Fischer, 1891, p._ 149). COCOS-KEELING IS-
LANDS: Direction Id. (ANSF; y); E. side of Horsburgh Id.;
SW side West Id. (both ANSP; y). CHRISTMAS ISLAND:
(Tomlin, 1934, p. 79). INDONESIA: Pulau Bai, Batu group,
off Sumatra; Pulau Stupai, Mentawai Ids., S.W. Sumatra;
Pelaboean Ratoe, Preanger, Java (all USNM; y); Morotai
Id., Moluccas (MCZ; y); Mantanani Id. and Mandi Darrah Id.,
N. Borneo (both ANSP; y); Batu Dua and Palau Pombo,
Wasi, Ambon I. (both FMNH; y); Malawali, N. Borneo (AMNH;
y). PHILIPPINES: Many localities throughout the Archi-
pelago; Luzon Id.; Mindoro Id.; Cebu Id.; Sulu Archipelago
(AMNH; ANSP; y); Samar Id.; Catanduanes Id.; Palawan Id.
(AMNH; ANSP; yw); Calamianes group; Mindanao Id.
AMNH: ANSP; MCZ; w); Borongan village, Samar Id.; Gig-
moto, Catanduanes Id. (both AIM; yw). FORMOSA (Taiwan):
Tainan beach (AMNH; w); Suo; Kasyo-to; Botanwan;
Ryukyu-syo; Hoko (Kuroda, 1941, p. 111; y); Karenko; Lasy-
oto; Garanbi; Hoko; Ryukyu-syo (Kuroda, 1941, p. 111; w).
RYUKYU ISLANDS: Kikaiga Shima (ANSP; y); Bolo reef,
NW Nakagami Gun, Okinawa (USNM; y); Okuma, Kuni-
gami-Gun; Odomari, Okinawa (both USNM; w); Yomitan
reef, Okinawa (AMNH; yw). JAPAN: Oshima, Osumi
(USNM; yw); Hachijo Id., off Honshu (ANSP; w); Tosa,
Shikoku (ANSP; y); Kagoshima, Kyushu (ANSP; w). MARI-
ANAS: Lagunan Tanapaa, Saipan; Agat Bay, Guam (both
ANSP; y); Apra Harbour, Guam (AMNH; y); Tinian Id.
(FMNH; yw); several localities on Guam Id. (USNM; w);
Saipan (AMNH; w). PALAU ISLANDS: Angupelu Id., SE
Koror Id.; reef N. of Gorokottan Id., S. side W. Passage,
Babelthuap Id. (both ANSP; y); S. E. of Auropushekaru Id.,
Malakal Harbour, Koror Id. (ANSP; yw); Angaur, Peleliu Id.
(FMNH; w). CAROLINE ISLANDS: N. of Kayangel Id.; S.
of Garakayo Id.; Ponape; Yap (all ANSP; y); Ulithi Atoll;
Satawal Atoll; Manini, Kapingamarangi (all USNM; y); S.E.
Rattakadokoru Id. (ANSP; w). MARSHALL ISLANDS: Eni-
wetok Atoll; Kwajalein Atoll (both AMNH; y); Bikini Atoll;
Eniwetok Atoll; Rongelap Atoll; Rongerik Atoll; Kwajalein
MicoBAR |:
MaLoy
13
ve et
ot © of mazamae's, TRICINUS RICINUS
160° 180° 160° igo
T T 7
90°
120° 100°
r
HEBRID
earn
| |
| @ YELLOW-SPOTTED FORM © 80TH FORMS
TEM Ap O WHITE FORM © UTERATURE RECORD l
—
|
60" ie ey = see T25*
Plate 17. Geographical distribution of Drupa (Drupa) ricinus
ricinus (Linnaeus) in the Indo-Pacific, and its subspecies
D. (D.) ricinus hadari Emerson and Cernohorsky, in the
Red Sea.
[14-826]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13
Drupa 23
Atoll (all ANSP; w). WAKE ISLAND: (AMNH, yw; ANSP,
y). NEW GUINEA: NE Noekori Id.; Manokwari; SW Biak
dock, Biak Id. (all ANSP; y); Boensaki Id., off Sowek, Soepi-
ori Id., Schouten Ids. (ANSP; yw); NE Mioes Woendi, Pad-
aido Ids. (ANSP; yw); Samarai, Papua (AMNH; yw); Huon
Gulf (USNM; w). AUSTRALIA: Queensland: Capricorn Ids.;
Hardy reef, Great Barrier reef (both AMNH; y); Birds Id.;
Green Id., Cairns (both USNM; y); Magnetic Id.; Lady Elliot
Id. (both AIM; y); New South Wales: (Iredale & McMichael,
1962, p. 74); West Australia: Vlaming Head (FMNH; y);
Houtman, Abrolhos Archipelago (Wilson & Gillett, 1971,
p. 92). NEW BRITAIN: Rabaul Harbour (ANSP; y). NEW
IRELAND: Kavieng (AMNH; y). SOLOMON ISLANDS:
Kieta, Bougainville; Lutee, Choiseul Id.; Ataa, N. Malaita
Id.; Tulagi Id., Santa Ana Id. (all AMNH; y); Tikopia Id.
(AIM; w); Lunga, Guadalcanal (AMNH; yw). NEW HEB-
RIDES: Lamap, Malekula Id. (ANSP; w); Bushmens Bay,
E. Malekula Id. (AIM; w); Pango Point, Efate Id. (AIM; w);
Tauna Id. (DM; w). NEW CALEDONIA: SE Dumbea Pass,
off Noumea; Baie Quemo, Noumea; Koe, Touho (all ANSP;
y). LOYALTY ISLANDS: Lifu (USNM; y). GILBERT IS-
LANDS: Abemama (USNM; w). ELLICE ISLANDS: Funa-
futi lagoon (USNM; AIM; w). WALLIS & FUTUNA IS-
LANDS: Faioa; Uvea; Nukuhifala, all Wallis Id. (USNM; y);
Nukuhifala, Wallis Id. (USNM; w); Anse de Sigave, Futuna
Id. (USNM; yw); Mua, Alofi, Hoorn Id. (USNM; y). FIJI
ISLANDS: Koro Bay reef, Vanua Levu (AMNH; y); Suva
Harbour, S. Viti Levu (USNM; y); Korolevu, S. Viti Levu;
Yasawa reef (both AMNH; w); Namagumagua village, S. Viti
Levu (WOC coll.; w); Rat Tail Passage, Suva reef, S. Viti
Levu; Mamanuca_ group; Caboni beach; Manava_ Id.;
Nananu-i-Ra Id.; Viti Levu Bay, all Viti Levu (all WOC coll.;
yw). TONGA ISLANDS: Ha’ateiho reef, Tongatapu; E. coast
Tongatapu Id. (both USNM; y); Haakoma, Tongatapu (MCZ;
w); Hufagalupe, Tongatapu (MCZ; yw) Niuafo’ou Id. (USNM;
w). NIUE ISLAND: near Malatu (USNM; y); Oneone reef
(AMNH; yw); Alofi (USNM; yw). SAMOA ISLANDS: Vailele
Bay, Upolu; Tafuna, Tutuila (both ANSP; y); Fagaitua Bay,
Tutuila; Ofu (both MCZ, w); Asau Harbor, Savaii (USNM;
yw); Apia, Upolu (AMNH; yw); Tau Id., Manua group (MCZ;
yw). TOKELAU ISLANDS: W. side Tukuo, Manihiki Atoll
(ANSP; w); Swains Id. (USNM; w). PHOENIX ISLANDS:
Canton Id.; Enderbury Id. (both USNM; w). HOWLAND IS-
LAND: (USNM; w). COOK ISLANDS: Bird’s Id., Tom’s Id.;
North Id., all Palmerston Atoll (all USNM; w); Koromiri Id.,
SE Rarotonga (ANSP; y); several localities on Aitutaki;
Mauke; Rarotonga; Mangaia (all MCZ; USNM; w); Motu
Akaiami, Aitutaki (USNM; yw). AUSTRAL ISLANDS:
Rurutu; Tubuai; Raivavae (all USNM; w). SOCIETY IS-
LANDS: Mopelia Id. (ANSP; y); many localities on Bora
Bora; Huahine; Raiatea; Moorea; Tahiti (USNM; w); Arue,
Tahiti; Taone, Tahiti; Huahine (all USNM; yw). TUAMOTU
ISLANDS: Napuka (AMNH; w); Tacume; Clermont Tenere;
Takaroa (all ANSP; w); Rangiroa (AMNH; yw); Mangareva
(USNM; w); Raroia (ANSP; yw). PITCAIRN ISLAND:
(AMNH; USNM; w). MARQUESAS ISLANDS: Taiohae,
Nukuhiva (AMNH; y); Hiva Oa Id. (ANSP; w); Ua Huka Id.
(AMNH; ANSP; w). LINE ISLANDS: Christmas Id.; Flint
Id.; Palmyra Id. (all ANSP; w); Palmyra Id. (MCZ; y); Jarvis
Id. (AMNH; y). JOHNSTON ISLAND: (USNM; w). HAWAI-
IAN ISLANDS: Honolulu Harbour (ANSP; y); Kure Id.;
Midway Id.; Laysan Id.; Tern Id.; French Frigate Shoal;
Kauai Id.; Oahu Id.; Molokai Id.; Maui Id.; Hawaii Id. (all
AMNH,; ANSP; USNM; w); Niihau Id.; Oahu Id. (both AIM;
w); Lihue, Kauai (AMNH; yw). CLIPPERTON ISLAND:
(AMNH,; ANSP; w); (Hertlein & Allison, 1960, p. 15; yw).
GALAPAGOS ISLANDS: (Hertlein, 1960, p. 8).
Fossil records— ZANZIBAR: Pleistocene: Base of well, vil-
lage W. of Makunduchi (Cox, 1927, p. 90, pl. 18, figs. 12a,
b and Stockley, 1928, p. 79). MOZAMBIQUE: Pleistocene:
Chidenguel, Inhambane district (Cox, 1939, p. 90). HAWAI-
IAN ISLANDS: Pleistocene: Oahu (Mansfield in Stearns and
Vaksvik, 1935, p. 167; Kosuge, 1969, p. 786, pl. 5, fig. 90).
Drupa ricinus new subspecies
hadari Emerson and Cernohorsky
(PI. 2, figs. 9, 10; Pl. 18)
Range—Red Sea only.
Remarks—Distinguished from the nominate
subspecies Drupa_ ricinus (Linnaeus) by _ its
larger size, heavier shell and more strongly de-
veloped parietal shield.
Habitat—Coral reefs and rocks, on intertidal
reef-flats.
Description—Juveniles typical of those of both
apertural color forms of the nominate subspecies,
with spinose nodules dark brown. Nodules on the
body whorl or mature specimens white, with
brown nodules on spire only, or body whorl ax-
ially streaked with brown except on the last two
or three rows of spines. Aperture in mature spec-
imens white with fully developed parietal shield
completely concealing nodular coloration of the
parietal area.
The side-cusps of the rachidian of the radula
are bifid and are followed by 2 weak, short lat-
eral denticles, excluding the slightly longer end-
cusps.
Measurements (mm)—(including spines)
length width
38.0 34.2 Eilat, Gulf of Aqaba (paratype
AMNH no. 112617b)
37.2 38.7 Eilat, Gulf of Aqaba (holotype
AMNH no. 166928)
34.2 34.1 Eilat, Gulf of Aqaba (paratype
AMNH no. 112617a)
32.0 32.0 Eilat, Gulf of Aqaba (paratype
AMNH no. 112617c)
28.0 29.0 Eilat, Gulf of Aqaba (paratype
DMNH no. 51119)
Synonymy—
1862 Ricinula albolabris Blainville, Ktister (pars), Syst.
Conchylien-Cabinet, ed. 2, Abt. IE, Ricinula, pl. 2,
figs. 1, 2 (Red Sea) [non Purpura albolabris Blain-
ville, 1832].
1941 Drupa (Drupa) ricinus (Linné), Wenz, Handb. Pali-
ozoologie, vol. 6, pt. 5, p. 1114, fig. 3165 (Red Sea)
{non Murex ricinus Linnaeus, 1758].
Pe pa Se
Can aaa
:
nN
“hy | J V/
Plate 18. Radula of Drupa (Drupa) ricinus hadari Emerson
and Cernohorsky. Half a transverse row; Eilat, Gulf of
Aqaba, Israel.
[14-827]
24 = Drupa
Types—The holotype, AMNH No. 166928, and
8 paratypes AMNH No. 112617, have been de-
posited in the American Museum of Natural His-
tory, New York. Other paratypes are in the Dela-
ware Museum of Natural History, Greenville,
The Auckland Institute and Museum and the Tel
Aviv University, Israel. The type locality is Eilat,
Gulf of Aqaba, Israel.
Nomenclature—The subspecies is named in
honor of the late Aryeh Hadar, who kindly sub-
mitted specimens from the Gulf of Aqaba for
study.
Records—RED SEA: Sharem, Gulf of Suez (DMNH): Eilat,
Gulf of Aqaba (AMNH; DMNH; AIM; w); Ras Banas, Egypt
(USNM; w); Quseir, Egypt (LACMNH; y); Mualla, Abu
Zabad, Gulf of Aqaba (Rees and Stuckley, 1952, p. 196);
Suakin, Sudan (Sturany, 1905, p. 141); Jiddah, Saudi Arabia
(AMNHII; yw); Pointe du Requin, [le Abulat (Franc, 1956,
p. 37).
Drupa elegans (Broderip and Sowerby, 1829)
(PI. 2, fig. 12; Pl. 19, 20)
Range—From Wake Island to the Tuamotu Is-
lands.
Remarks—This species is similar to Drupa ri-
cinus (Linnaeus), but differs in apertural color-
ation, the lack of tubercle coloring and in being
smaller in size. This uncommon species occurs
sympatrically throughout its range with D. ri-
cinus (Linnaeus). It may prove eventually to be
another, moderately rare color phase of that
variable species. Although an occurrence of
D. elegans in the Society Islands is probable, the
cited record requires confirmation; specimens
from that locality in the Academy of Natural
Sciences of Philadelphia and the National Mu-
seum of Natural History, Washington, lack exact
locality data.
Habitat—Only five specimens were taken by the
operation “Crossroads” in 1946 and the biologi-
cal re-survey in 1947 of the Marshall Islands.
These were found living in association with the
extremely common _ yellow-spotted and_ white
color phases of Drupa ricinus. On Niue Island,
the species is also sympatric with D. ricinus and
has been collected on reefs under cliff overhangs
(D. C. Johnson, personal commun.).
Description—Shell 13 to 25 mm (about % to 1
inch) in length, sub-ovate, spire short; body
whorl spirally ribbed with five rows of siphonous
Emerson and Cernohorsky
Muricidae
Limam
Plate 19. Radula of Drupa (Drupa) elegans (Broderip and
Sowerby). Half a transverse row; Vailoa, Alofi Bay, Niue
Id.
Plate 20. Drupa (Drupa) elegans (Broderip and Sowerby).
Figs. 1, 2. Lectotype from unknown locality (B.M. (N.H.),
20.3 x 22.4 mm) [photo courtesy J. Taylor, B. M. (NH)].
Figs. 3, 4. Specimen from Vailoa, Alofi Bay, Niue Id.
(WOC coll.; 19.0 x 20.7 mm).
tubercles. Intersticial surface minutely scaled.
Parietal shield moderately enameled, reflected
over body wall to form a heavy callus; lower por-
tion of columella 4 plaited. Axial fold strongly
developed. Aperture very narrow, dentition of
[14-828]
May 30, 1973
outer lip as in Drupa ricinus (Linnaeus). Color of
the exterior and aperture white; aperture encir-
cled by a continuous dark riddish brown line in
mature specimens; in juvenile specimens, the line
is disrupted and restricted to the columellar lip
and near the anal siphonal canal.
Radular ribbon very small, side-cusps of the
rachidian bifid, followed by 3. slender and
deeply rooted lateral denticles, exclusive of the
short end-cusps.
Measurements (mm)—(including spines; all
specimens with a mature lip)
length width
25.0 25.0 Wake Island
20.3 22.3 Lectotype of elegans B. and S.
19.0 20.7 Niue Id.
18.5 17.5 Lord Hood Id. (S. Marutea Id.)
13.6 11.8 Lord Hood Id. (S. Marutea Id.)
Synonymy—
1829 Ricinula elegans Broderip & Sowerby, Zoological Jour-
nal, London, vol. 4, p. 376 (no locality given); 1839
Gray, Zool. Capt. Beechey’s voyage, p. 155, pl. 36,
fig. 4; 1844 Deshayes & Milne-Edwards, Hist. nat.
anim. s. verttbres, ed. 2, vol. 10, p..52; 1846 Reeve,
Conchologia Iconica, vol. 3, pl. 1, fig. 1 (Lord Hood
Id. = South Marutea Id.); 1933 Dautzenberg & Bouge,
Journal de Conchyliologie, vol. 77, p. 238.
1853 Pentadactylus elegans Broderip, H. & A. Adams, Gen-
era Recent Mollusca, vol. 1, pp. 129, 130; vol. 3, pl.
13, figs. 6a, b (operculum only).
1880 Ricinula ricinus var. elegans Broderip, Tryon, Manual
of Conchology, vol. 2, p. 184, pl. 56, fig. 193.
1913 Drupa ricinus var. elegans Broderip & Sowerby, Hed-
ley, Nautilus, vol. 27, no. 7, p. 80.
1957 Drupa elegans Broderip & Sowerby, Kaicher, Indo-
Pacific Sea Shells, (Muricacea, Buccinacea), pl. 3,
fig. 18.
Types—Three syntypes of Drupa elegans are in
the British Museum (Nat. Hist.), and the 20.3mm
long syntype (PI. 20, figs. 1, 2) is here selected
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa = 25
as the lectotype. The species was described from
shells in the Museum of the Zoological Society
brought home by Lieutenant Belcher who sailed
under Captain Beechey in the “Blossom” on its
voyage to the Bering Straits and the Pacific
(1825-1828). No type locality was given, but
Cuming collected it from Lord Hood Island (=
S. Marutea Id.), and his specimen was figured
by Reeve (1846). Lord Hood Island (= S. Marutea
Id.) is here designated as the type locality.
Records—WAKE ISLAND: (BPBM). MARSHALL IS-
LANDS: Namu Id., N. W. end of Bikini Id.; Eniman Id.,
Bikini Atoll; Eniwetok Atoll (all USNM). LINE ISLANDS:
Caroline Id. Flint Id. (both ANSP). NIUE ISLAND: (AMNH);
Vailoa, Alofi Bay (D. C. Johnson coll.; WOC coll.); Avatele
(S. Herriot coll.). COOK ISLANDS: Akamaru Id., Manihiki
Atoll (ANSP). SOCIETY ISLANDS: (ANSP; USNM). TU-
AMOTU ISLANDS: Lord Hood Id. (= S. Marutea_ Id.)
(AMNH; BMNH); Vahitahi; Nengonengo Id. (both USNM);
Anaa Id. (MCZ); Napuka; Taenga; Fakahina (Dautzenberg
& Bouge, 1933, p. 238).
140° 160° 180° 160° 140° 120°
CAROLINE
wove TELEGANS
6.4 —_| =
NEW
ZEALAND
Sp |
1
160° 180°
Plate 21. Geographical distribution of Drupa (Drupa) ele-
gans (Broderip and Sowerby). Open circles are literature
records.
[14-829]
26 Drupa Emerson and Cernohorsky Muricidae
ps ae ce en ie Ae
[These occasional blank areas occur between genera and subgenera to permit
the insertion of new material and future sections in their proper systematic
sequence. }
[14-830]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa = 27
Subgenus Ricinella Schumacher, 1817
Type: Drupa rubusidaeus Réding, 1798
Shell sub-ovate, solid, spire rather short; body
whorl large, whorls spirally ribbed, sculptured
with siphonous spiniform tubercles; interstices
filled with fine scale-like plates forming raised
threads; threads running spirally in close parallel
association; columella with an axial fold, outer
lip dentate with teeth singularly arranged, some-
times in close association, not compound; oper-
culum subelliptical, not linear; radula typical for
the genus.
This group includes besides the type species,
Ricinula speciosa Dunker, 1867, R. clathrata
clathrata Lamarck, 1816, and R. clathrata miti-
cula Lamarck, 1822. On the basis of radular
characters, the group shows a close _relation-
ship with Drupa sensu stricto. The shells on the
other hand resemble some of the thaids. The
members of this group have the labial teeth de-
veloped as small, pearl-like teeth which may be
closely crowded together as in Drupa clathrata
or D. speciosa, but are never actually compound.
The interpretation of the identity of the type
species of Ricinella, i.e. R. purpurata Schu-
macher, rests on the elucidation of the cited illus-
tration of Favanne (1784, pl. 24, fig. 2). This fig-
ure resembles D. rubusidaeus Riding more so
than D. clathrata (Lamarck). Favanne’s descrip-
tion of the color of the aperture as being deep
and vidid lilac or purple, excludes the species
D. clathrata from consideration.
Synonymy—
1817 Ricinella Schumacher, Essai nouv. systéme, pp. 72,
240. Type-species by subsequent designation, Iredale,
1937: Ricinella| purpurata Schumacher, 1817 [=
Drupa rubusidaeus Réding, 1798].
| pa
Plate 22. Radula of Drupa (Ricinella) rubusidaeus Réding.
Half a transverse row; Olawala, Maui, Hawaiian Ids.
Drupa rubusidaeus Réding, 1798
(Pl. 2, figs. 13-15; Pls. 22, 23)
Range—From the Red Sea and East Africa
throughout the Indo-Pacific to Hawaii and the
Tuamotu Islands.
Remarks—This solid species could only be con-
fused with the much smaller and rarer Drupa
speciosa (Dunker). D. speciosa is a higher-spired
shell with no yellow coloration in the aperture;
the pink coloring of the aperture is more pro-
nounced and the denticles of the outer lip are
more closely set and almost grouped.
Habitat-—On algae matted reef-flats, under
rocks and in tide-pools, generally on the seaward
half of the reef-flat; from low tide to a depth of
10 fathoms or more, on rock, sand or coral. In
the Hawaiian Islands the species has been col-
lected attached to stony coral at a depth of 50 to
60 feet (C. S. Weaver, leg.).
Description—Shell 20 to 55 mm (% to 2% inches)
in length, solid, ovately globose, slightly ventri-
cose, spire short, acuminated, body whorl large
and with five rows of strong, sub-spiniform,
siphonous tubercles. Sculpture of fine, scale like
Plate 23. Drupa (Ricinella) rubusidaeus Réding.
Fig. 1. Lectotype figure of D. (R.) rubusidaeus Rodding,
from Knorr, 1768, pt. 6, pl. 24, fig. 7.
Fig. 2. Holotype of Ricinula reeveana Crosse, from Nuku-
hiva, Marquesa Ids. (B.M. (NH); 48.2 x 38.8 mm—im-
mature specimen).
[14-841]
28 Ricinella
plates forming ridges which run spirally in par-
allel association between the rows of tubercles.
Aperture sub-elliptical, outer lip in adult stage
dentate with 7-12 small pearl-like teeth; margin
crenulated, apertural area enameled; enameled
area extending over a large portion of the body
whorl adjacent to the aperture. Inner lip cal-
lused with 3 or 4 plications. Siphonal canal
short, deep, recurved; sutural canal conspicuous,
always open. Color white-yellowish externally,
margin of the aperture yellow, interior of the
aperture rich pink.
The side-cusps of the rachidians of the radula
are bifid to trifid and are followed by 2-3 lateral
denticles, exclusive of the stronger side-cusps.
Measurements (mm)—(including — spines; all
specimens with a mature lip)
length width
53.8 48.5 Oahu, Hawaiian Ids.
48.2 38.8 Holotype of reeveana Crosse
41.6 38.9 Luzon Id., Philippines
34.0 28.7 Niue Id., Polynesia
28.8 25.0 Koror Id., Palau Ids.
Synonymy—
1742 Gualtieri, Ind. test. Conchyliorum, pl. 28, fig. R
(poor).
Regenfuss, Aus. Schnecken, Muscheln u.a.Schaal-
thiere, (1), pl. 3, fig. 32.
Knorr, Verg. Augen U. Gemiiths, pt. 6, pl. 24,
fig. 7 (very good).
1777 “Murex hericinus” Martini, Syst. Conchylien-Cabinet,
vol. 3, p. 283, pl. 101, figs. 974, 975 (poor). [non-
binomial].
Favanne, La Conchyliologie
d’Argenville], pl. 24, fig. A2 (poor).
Favanne, Catalogue systematique et raisonné
ee ates p. 145 (description).
1791 Murex nodus Gmelin, Systema Naturae, ed. 13, p. 3537
(refers to Knorr, op. cit., with a query ) [no locality
given] (non Linnaeus, 1758).
1798 Drupa rubusidaeus Roéding, Museum Boltenianum,
p. 55 (refers to Martini, op. cit., and Knorr, op. cit.)
[no locality given]; 1913 Hedley, Nautilus, vol. 27,
no. 7, p. 80; 1966 Weaver, Hawaiian Shell News,
vol. 14, no. 14, p. 2, textfigs. 1, 2; 1969 Cernohorsky,
Veliger, vol. 11, no. 4, p. 301, pl. 47, figs. 10, 10a;
1971 Wilson & Gillet, Australian Shells, p. 92, pl. 61,
figs. 1, la. ,
1798 Drupa fragum hoding, Museum Boltenianum, p. 55
(refers to Murex nodus Gmelin, 1791, sp. 42) [no
locality given].
1807 Mancinella hystrix Link, Beschr. Nat.-Samml. Univ.
Rostock, 3 Abth., p. 115 (refers to Martini, op. cit.)
[non Murex hystrix Linnaeus, 1758].
1817 Ricinella purpurata Schumacher, Essai nouv. systéme,
p. 240 (refers to Favanne, op. cit.) [no locality given].
1817 Murex hystrix Linnaeus, Dillwyn, Desc. cat. Rec. shells,
vol. 2, p. 706 (refers to Martini, Gualtieri, Regenfuss
etc.) [East Indian Seas; coasts of the Friendly Islands
= Tonga Ids.]; 1825 Wood, Index Testaceologicus,
p. 124, pl. 26, fig. 50a (non M. hystrix Linnaeus,
1758).
1758
1768
1780
ed. 3. [of
1784
Emerson and Cernohorsky
Muricidae
1822 Purpura hystrix Lamarck, Hist. nat. anim. s. vertebres,
vol. 7, p. 247 (refers to Knorr, Regenfuss, Martini and
Gualtieri, op. cit.); 1835 Kiener, Spéc. gén. icon. coq.
viv., vol. 8, p. 13, pl. 2, figs. 4, 4a, b; 1846 Reeve,
Conchologia Iconica, vol. 3, pl. 3, fig. 13 (non Murex
hystrix Linnaeus, 1758).
1825 Murex hippocastanum Wood, Index Testaceologicus,
p. 124, pl. 26, fig. 53a (non Linnaeus, 1758).
1832 Purpura spathulifera Blainville, Nouv. Ann. Mus.
(Hist. Nat. Paris, ser. 3, vol. 1, p. 212, pl. 9, fig. 8
(no locality given).
1833 Purpura histrix (sic) Lamarck, Quoy & Gaimard, Voy-
age L’Astrolabe, vol. 2, p. 575, pl. 39, figs. 14-16
(animal and operculum).
1853 Pentadactylus hystrix H. & A. Adams, Gen. Rec. Mol-
lusca, vol. 1, p. 130; 1875 Troschel, Gebiss d.
Schnecken, vol. 2, p. 134, pl. 13, fig. 4 (radula).
1862 Ricinula reeveana Crosse, Journal de Conchyliologie,
vol. 10, p. 47, pl. 1, fig. 3 (Nouhiva = Nukuhiva, Mar-
quesas Ids.) [immature specimen] (non Ricinula
reeveana C. B. Adams, 1852).
1880 Ricinula hystrix Linné, Tryon, Manual Conchology, vol.
2, p. 183, pl. 56, tig. 195; 1933 Dautzenberg & Bouge,
Journal de Conchyliologie, vol. 77, p. 239.
1880 Ricinula hystrix var. reeveana Crosse, Tryon, ibid.,
vol. 2, p. 183, pl. 56, fig. 196; 1933 Dautzenberg &
Bouge, ibid., vol. 77, p. 239 (non C. B. Adams, 1852).
1911 Sistrum hystrix Linné, Schepman, Siboga-Expeditie,
vol. 49d, p. 354 (non Murex hystrix Linnaeus, 1758).
1913 Drupa rubusidaeus Bolten, Hedley (pars), The Nau-
tilus, vol. 27, no. 7, p. 79.
1936 Drupa spathulifera (Blainville), Hirase, Coll. Jap. shells,
ed. 5, p. 79, pl. 110, fig. 9; 1954 Kira, Col. illust.
shells Japan, p. 47, pl. 23, fig. 4; 1967 Habe & Ko-
suge, Stand. book Jap. shells color, vol. 3, p. 70,
pl. 27, fig. 24; 1968 Taylor, Phil. Trans. Roy. Soc.
London, ser. B, vol. 254, p. 201.
1938 Drupa (Drupa) hystrix (Linné), Adam & Leloup, Mém.
Mus. Roy. d’Hist. Nat. Belgique, vol. 2, fasc. 19,
p. 163.
1957 Drupa rubuscaesia Kaicher, Indo-Pacific Sea Shells
(Muricacea, Buccinacea), pl. 3, fig. 15 (non D. rubus-
caesius Réding, 1798).
1961 Drupa rubuscaesium Rodding, Rippingale & McMichael,
Queensland and Gt. Barrier reef Shells, p. 102, pl. 13,
fig. 3 (non D. rubuscaesius Roding, 1798).
1962 Drupa rubuscaesius (R6éding), Kira, Shells West. Pa-
cific in color, vol. 1, p. 62, pl. 24, fig. 4; 1965 Ara-
kawa, Venus: Jap. Journ. Malacology, vol. 24, no. 2,
p. 115, pl. 13, figs. 5, 6 (radula) [non Réding, 1798].
1965 Drupa speciosa (Dunker), Wu, Bull. Inst. Zool. Acad.
Sinica, vol. 4, p. 98, textfig. 30 (radula) [non Ricin-
ula speciosa Dunker, 1867].
1967 Drupa rubusidaea Réding, Orr Maes, Proc. Acad. Nat.
Sci. Philadelphia, vol. 119, no. 4, p. 129.
T ypes—Réding’s type specimen of Drupa rubu-
sidaeus is lost, and we therefore select the spec-
imen figured by Knorr on plate 24, fig. 7, as the
lectotype of the species (Pl. 23, fig. 1), and
designate Davao Bay, Mindanao, Philippines, as
the type locality. The type specimen of Purpura
spathulifera Blainville, is probably in the Mu-
séum National d’Histoire Naturelle, Paris, and
the holotype of Ricinula reeveana Crosse, is in
the British Museum (Nat. Hist.) [Pl. 23, fig. 2].
The type specimen of Ricinella purpurata Schu-
[14-842]
May 30, 1973
o9°
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13
Drupa_ 29
20°|—
ees:
or
RUBUSIDAEUS] ‘y
CHAOS
s.?
Plate 24. Geographical distribution of Drupa (Ricinella) rubu-
sidaeus Réding. Open circles are literature records.
macher, could not be located in the Zoological
Museum, Copenhagen (J. Knudsen, in litt.).
Nomenclature—This species was at first mas-
querading under the names Mancinella hystrix
Link, Murex hystrix Dillwyn and Purpura hystrix
Lamarck. As previously discussed, this is not
Murex hystrix Linnaeus, 1758. Since Réding,
1798, cites 2 of the 4 references which Lamarck
considered synonyms of Purpura hystrix, there
can be no doubt regarding the identity of his
species. Réding’s citation to Knorr’s excellent
figure (designated as lectotype) of the present
species, as well as Martini’s figures, serves to
establish the identity of D. rubusidaeus.
More recently Hedley (1913) confused Purpura
spathulifera Blainville, a species which he synon-
ymized with Drupa rubuscaesius Roding, Ricin-
ula clathrata Lamarck, and R. speciosa Dunker,
but which is conspecific with D. rubusidaeus.
The specimen and its figured radula cited by Wu
(1965a) as “Drupa speciosa (Dunker)” originated
from Wan-li-tong, Taiwan, and is actually D.
rubusidaeus Réding (Wu in litt.).
Records—RED SEA: (AMNH). EAST AFRICA: Gesira,
Somalia (ANSP); Diani Beach, Kenya (Heinicke, 1970, p. 7);
Kendwa Id., 4 mi. ESE of Dar-es-Salaam; Sinda Id., 15 mi.
SSE of Dar-es-Salaam; N.W. of Magogani; Pangavini Id., 10
mi NNE of Dar-es-Salaam, all Tanzania (all MCZ); Mozam-
bique (USNM). SEYCHELLES ISLANDS: Beau Vallon
Beach, Mahe; Cousin Id. (both ANSP). MADAGASCAR:
Nosy N’Tangam, W. Nossi Bé; Grande Recife, W. of airport,
Tulear; Ambodifototra (all MCZ). REUNION ISLAND:
(ANSP). MALDIVE ISLANDS: Miladummadula Atoll; Tila-
dummati Atoll; N. Malé Atoll (all ANSP). LACCADIVE
ISLANDS: (Hornell, 1922, p. 217). CEYLON: (Langdon,
1875, p. 72). VIETNAM: Cam Ranh Bay (ANSP). COCOS-
KEELING ISLANDS: Klapetuju, West Id. (USNM); S. end of
Home Id.; N. end of Horsburgh Id.; S. end of Direction Id.
(all ANSP). CHRISTMAS ISLAND: (Tomlin, 1935, p. 79).
INDONESIA: Mandi Darrah Id., N. Borneo (AMNH;
ANSP); Pulau Bai, Batu group, off Sumatra; Pulau Penju,
S. Sumatra (both USNM); Bali Id.; Wasi, Ambon Id. (both
FMNH); Banda Id.; Soengai Manoembaii, Hes Aroe (Adam
& Leloup, 1938, p. 164); Tjilaoet, Java (Altena, 1945,
p. 146). PHILIPPINES: Iba, Zambales, Luzon Id.; Boron-
gan village, E. Samar Id.; Cuyo Id., Palawan group (all
ANSP); Tilig reef, Lubang Id.; Calapan, Mindoro: Linga-
yan Gulf; Davao Bay, Mindanao (all AMNH); Nogas Point,
Panay Id.; Cabra Id.; Silino Id.; Point Matangal, Basilan,
Sulu (all USNM). FORMOSA (Taiwan): Wan-li-tong (AMNH);
Karenko; Botan-wan; Ryuku-syo; Hoko (Kuroda, 1941,
p. Ill). RYUKYU ISLANDS: Yomitan’ reef, Okinawa
(AMNH); Okuma, Kunigami-Gun; Bolo reef, N.W. Naka-
gami-Gun (both USNM); Ishigake (MCZ); Miyako (FMNH).
JAPAN: Hachijo Id., 275 mi. S. of Tokyo; Tosa, Shikoku;
Kagoshima, Kyushu (all ANSP); Ominato Ise (AIM); Oshima,
Osumi (Powell coll.). MARIANAS: Saipan (AMNH); Agana
Bay, Guam Id. (ANSP). PALAU ISLANDS: Babelthuap Id.;
Angupelu Id., S.E. of Koror; Malakal Harbour, Koror; Helen
Channel (all ANSP). CAROLINE ISLANDS: Kayangel;
Ngargersiul; S.W. of Rattakadokoru; S. of Garakayo; Ulithi;
Ella; Elangalap Id., Ifaluk Atoll; Tirakaume; Ringutoru;
Kapingamarangi (all ANSP); Ponape reef (AMNH). WAKE
ISLAND: (BPBM). MARSHALL ISLANDS: Eniwetok Atoll,
Bikini Atoll; Rongerik Atoll (all USNM); Kwajalein Atoll
(Dietrich & Morris, 1953, p. 15). NEW GUINEA: Biak Id.,
Schouten Ids. (USNM); 1 mi. NE of Mioes Woendi, Padaido
Ids. (Powell coll.); Wewak (ANSP). NEW BRITAIN: Rabaul
Harbour (AMNH). AUSTRALIA: Queensland: Herald Cay,
Coral Sea; Holmes reef, Coral Sea; off Cairns; Watt reef, off
Townsville (all AMS); Green Id., off Cairns (AIM); several
islands of the Capricorn group (AMS; AMNH). SOLOMON
ISLANDS: Choiseul Bay, Choiseul Id.; Bougainville Id.;
Bellona Id. (all AMNH); Fiu, Malaita Id. (Powell coll.). NEW
HEBRIDES: Efate Id. (Colardeau coll.); Espiritu) Santo
(Solem, 1959, p. 262). NEW CALEDONIA: Touho (AMNH).
GILBERT ISLANDS: Abaiang (MCZ); Kingsmill Ids.
(AMNH). ELLICE ISLANDS: Funafuti lagoon (AIM). FIJI
ISLANDS: (AMNH,; ANSP); Rat Tail Passage, Suva reef,
S. Viti Levu (WOC coll.); Mamanuca group (Jennings coll.).
NIUE ISLAND: (DM); Tuapa reef (AMNH; WOC coll.).
SAMOA ISLANDS: reef at Satalo Id., Upolu Id.; E. side of
Wailele Bay, Upolu Id. (both ANSP); Nuuli, Tutuila Id.
(MCZ). COOK ISLANDS: Mauke; Aitutaki (both USNM);
off Aroa Creek, S.W. Rarotonga; Koromiri Id., S.W. Raro-
tonga (both MCZ); Mangaia (DM). SOCIETY ISLANDS:
Several localities on Tahiti and Moorea (USNM). TUA-
MOTU ISLANDS: Anaa Id. (AMNH; ANSP; MCZ); Amanu
(Coututirer, 1907, p. 143). MARQUESAS ISLANDS: (MCZ);
Nukuhiva (BMNH). LINE ISLANDS: Fanning Id. (DMNH).
HAWAIIAN ISLANDS: Midway Id.; off Waikiki, Oahu (both
[14-843]
30 Ricinella
Emerson and Cernohorsky
Muricidae
AMNH); Makaha Point, $.W. Oahu (MCZ); Olawala, Maui
(AMNH).
Fossil records—KENYA: Pleistocene: raised reef, S.E. of
Mombassa Id., N. of Ras Serani (Cox, 1930, p. 145). HA-
WAITAN ISLANDS: Oahu Id. (Kosuge, 1969, p. 786, pl. 5,
fig. 94).
Drupa speciosa (Dunker, 1867)
(Pl. 2, figs. 21, 22; Pl. 25)
Range—Tuamotu and Pitcairn Islands.
Remarks—The shell of this Polynesian species
is smaller and higher-spired than D. rubusidaeus
Réding, with a deeper pink to mauve aperture
which lacks the yellow coloration found in the
aperture of the larger species. Specimens labelled
“Fiji Islands” are obviously based on erroneous
locality indications as the species does not occur
there. The record from Rarotonga, Cook Islands
(ANSP) is also suspect, and has not been con-
firmed by recently collected, well-documented
specimens.
Habitat—Unknown, but probably found on
coral reefs at low tide.
Description—Shell 20 to 29 mm (% to 14 inches)
in length, solid, ovate, globose; spire moderately
elevated, acuminate; body whorl axially ribbed
with 9 ribs, crossed by 5 transverse rows of
short, strong, spinose tubercles. Interstices be-
tween tubercles striated with parallel rows of
small scales. Aperture subelliptical, columella
posteriorly excavated, outer lip dentate with 4-7
evenly spaced small white teeth; margin crenu-
lated. Inner lip with 3 strong lower plications and
1 subobsolete upper plication. Apertural area
enameled, enameled area of the inner lip extend-
ing over a portion of the body whorl. Anterior
siphonal canal short, deep; posterior siphonal
canal open. Color creamy-white externally, aper-
ture a deep mauve pink.
Radula unknown. The radular dentition figured
by Wu (1965a) for this species was based on a
specimen of D. rubusidaeus from Taiwan (Wu,
in litt.).
Measurements (mm)—(including spines; all
specimens with a mature lip)
length width
27.0 21.5 Tuamotu Ids.
26.9 23.2 Lectotype of speciosa Dunker
26.4 20.8 Raroia, Tuamotu Ids.
25.4 21.7 Vahitahi, Tuamotu Ids.
22.8 18.9 Vahitahi, Tuamotu Ids.
21.9 16.3 Vahitahi, Tuamotu Ids.
20.1 14.8 Tuamotu Ids.
Synonymy—
1846 Ricinula clathrata Lamarck, var. B., Reeve, Conchol-
ogia Iconica, vol. 3, pl. 2, fig. 9a (Anaa Id.) [non R.
clathrata Lamarck, 1816].
1867 Ricinula speciosa Dunker, Novitates Conchologicae,
Abt. I, pts. 11/12, p. 100, pl. 33, figs. 7, 8; 1878
Dunker, Addenda & Corrigenda, p. 139 (Philippine
Ids. = error) [non Purpura speciosa Valenciennes,
1832].
1880 Ricinula hystrix var. speciosa Dunker, Tryon, Manual
Conchology, vol. 2, p. 183, pl. 56, fig. 194; 1933
Dautzenberg & Bouge, Journal de Conchyliologie, vol.
77, p. 239.
1969 Drupa speciosa Dunker, Cernohorsky, Veliger, vol. 11,
no. 4, pp. 301, 302.
Types—Dunker described the species from 4
specimens in Hugh Cuming’s collection which is
now in the British Museum (Nat. Hist.). Three of
the syntypes are immature examples with weak
denticles on the outer lip and a superficial col-
umellar callus. Only one specimen is reasonably
mature, and this specimen is here selected as the
lectotype of Ricinula speciosa Dunker (P1 25, fig.
1). Even though the selected lectotype is without
Plate 25. Drupa (Ricinella) speciosa (Dunker).
Fig. 1. Lectotype (B.M. (NH); 26.9 x 23.2 mm).
Fig. 2. Syntype; immature (B.M. (NH); 26.9 x 21.9 mm)
{photos courtesy K. Way, B.M. (NH) ].
Fig. 3. Specimen from Vahitahi, Tuamotu Ids. (USNM
613343; 22.8 x 19.0 mm).
[14-844]
May 30, 1973
question the Drupa speciosa of authors, the spec-
imen does not compare too well with Dunker’s
original type-figure. Either the artist exercised
his own imagination when depicting the spec-
imen, or the originally illustrated example has gone
astray. Stability of the taxon Drupa speciosa,
however, is best served by the selection of the
most mature specimen from among the 4 syn-
types, which all are the D. speciosa of Dunker
and of authors. Dunker gave the erroneous type
locality as “Philippine Islands,” which is here
corrected to Anaa Island, Tuamotu Islands.
Reeve’s figure is based on a specimen in the
Cuming collection from this locality.
Nomenclature—Reeve in 1846 figured this spe-
cies, calling it Ricinula clathrata Lamarck, vari-
ety B, and both Crosse (1862) and Dunker (1867)
in describing reeveana and speciosa respectively,
cite Reeve’s figures in synonymy. However, R.
reeveana, which is preoccupied (non C. B. Ad-
ams, 1852), has been shown to be a synonym of
D. rubusidaeus Réding, leaving Dunker’s name
available for the present taxon.
Records—TUAMOTU ISLANDS: (USNM; ANSP; FMNH;
AMS); Garumaoa; Mataira; Oneroa; Opakea, all Raroia Atoll
(USNM); Anaa Id. (MCZ); Vahitahi (USNM); Fangatau;
Makatea (Dautzenberg and Bouge, 1933, p. 239). PITCAIRN
ISLAND: Oeno Island (USNM).
+ Pearl f Hermes
LL Is.
iA 3 Christmes
. es
an ©
= ‘PHoEN 1s. .
+ TOKELAU IS. 4 + MARQUESAS IS.
Same, “.
v “830A ts
NEW .
gee + Moria Theresa
tS (as rn ae
Plate 26. Geographical distribution of Drupa (Ricinella)
speciosa (Dunker). Open circles are literature records.
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 31
Drupa clathrata clathrata (Lamarck, 1816)
(Pl. 2, figs. 16-18; Pls. 27, 28)
Range—From Japan and the Philippines to the
Marquesas and Pitcairn Islands.
Remarks—The large number of spines, rugose
columella and apertural coloration serve to char-
acterize this species. Populations occurring in the
Indian Ocean are separable on shell morphology
and are'recognized as a subspecies, D. clathrata
miticula (Lamarck).
Habitat—On reefs, in crevices and under coral
rocks, intertidal.
Description—Shell 16 to 57 mm (% to 2% inches)
in length, ovate and solid, spire short; body
whorl large, crossed by five rows of spiral ribs.
Ribs with siphonous, spiny tubercles; tubercles
higher towards the margin of the outer lip. Spiral
ribs connected by low axial ribs to form shallow
pockets in the interstices. Surface sculptured
with fine spirally arranged scales. Aperture sub-
elliptical, columella excavated posteriorly, with
4 to 5 small plications above the siphonal canal.
Parietal shield thinly enameled, reflected to form
an irregular callus. Sutural canal well developed
as a groove turned toward the spire. Axial fold
strong, extending the length of the columella and
terminating as a margin of the siphonal canal.
Outer lip dentate with 4 to 5 singularly arranged
teeth which sometimes become united. Color
brownish-white externally, margin of aperture
spotted with brown, interior of aperture a light
violet. Operculum typical of group.
The radula has the side-cusps of the rachidian
bifid or trifid, and there are 2 to 3 deeply rooted
lateral denticles, exclusive of the end-cusps.
O.1MM =I
Plate 27. Radula of Drupa (Ricinella) clathrata clathrata
(Lamarck). Half a transverse row; Pango Point, Efate Id.,
New Hebrides.
[14-845]
32. Ricinella
Plate 28. Figs. 1, 2. Drupa (Ricinella) clathrata clathrata
(Lamarck); Fig. 3. D. (R.) clathrata miticula (Lamarck).
Fig. 1. Holotype of Ricinula clathrata Lamarck (MHNG
no. 1101/14/1; 30.4 x 31.2 mm).
Fig. 2. Specimen of Drupa (Ricinella) clathrata clathrata
(Lamarck), from Pango Point, Efate Id., New Hebrides
(WOC coll.; 28.3 x 25.8 mm).
Fig. 3. Lectotype of Ricinula miticula Lamarck (MHNG no.
1101/13/1; 26.3 x 21.0 mm).
Measurements (mm)—(including
specimens with a mature lip)
spines; all
length width
57.3 53.0 “South Seas”
42.8 38.2 “Central Pacific”
30:1 29.0 Suva reef, Fiji Ids.
32.2 24.6 Samar Id., Philippines
30.4 31.2 Holotype of clathrata Lamarck
23.1 20.0 Tacume Id., Tuamotu Ids.
21.0 18.5 Pango Pt., New Hebrides
Synonymy—
1816 Ricinula clathrata Lamarck, Tableau Encycl. Méthod-
ique, p. 2, pl. 395, figs. 5a, b (no locality given);
1822 Lamarck, Hist. nat. anim. s. vertébres, vol. 7,
p. 231; 1846 Reeve, Conchologia Iconica, vol. 3, pl. 2,
fig. 9b only (Elizabeth Id., = Tuamotu Ids.); 1859
Chenu, Manuel de Conchyliologie, vol. 1, p. 168,
textfig. 816; 1933 Dautzenberg & Bouge, Journal de
Conchyliologie, vol. 77, p. 237.
Emerson and Cernohorsky
Muricidae
1835 Purpura clathrata Lamarck, Kiener (pars), Spéc. gén.
icon. coq. viv., vol. 8, p. 15, pl. 3, fig. 5 only.
1853 Pentadactylus clathratus Lamarck, H. & A. Adams,
Gen. Rec. Mollusca, vol. 1, p. 130; 1875 Troschel,
Gebiss d. Schnecken, vol. 2, p. 133, pl. 13, fig. 3
(radula).
1880 Ricinula hystrix var. clathrata Lamarck, Tryon, Manual
Conchology, vol. 2, p. 184, pl. 56, figs. 197, 198.
1913 Drupa rubuscaesia Bolten, Hedley (pars), Nautilus,
vol. 27, no. 7, p. 80 (non D. rubuscaesius Réding,
1798).
1936 Drupa rubuscaesia Réding, Hirase, Coll. Jap. shells,
p. 79, pl. 110, fig. 10; 1957 Kaicher, Indo-Pacific Sea
Shells (Muricacea, Buccinacea), pl. 3, fig. 15 (non
D. rubuscaesius Réding, 1798).
1951 Drupa rubuscaesius Réding, Hirase & Taki, Handb.
illust. shells colour, pl. 110, fig. 10; 1959 Kira, Col.
illust. shells of Japan, vol. 1, p. 58, pl. 23, fig. 9
(non Réding, 1798).
1954 Drupa rubuscaesium Réding, Kira, Col. Ilust. shells
of Japan, pl. 23, fig. 9 (non D. rubuscaesius Réding
1798).
1962 Drupa rubsidaeus (sic) Réding, Kira, Shells west. Pa-
cific in colour, p. 63, pl. 24, fig. 9 (non D. rubusid-
aeus Réding, 1798).
1965 Drupa rubusidaeus Roding, Arakawa, Venus: Jap.
Journ. Malacology, vol. 24, no. 2, p. 115, pl. 13, fig. 7
(radula) [non Réding, 1798].
1967 Drupa_ (Ricinella) rubusidaeus (Roéding), Habe &
Kosuge, Stand. book Jap. shells in color, vol. 3, p. 70,
pl. 27, fig. 29 (non Roding, 1798).
1969 Drupa clathrata (Lamarck), Cernohorsky, Veliger, vol.
11, no. 4, p. 298, pl. 47, fig. 6.
Types—The holotype of Ricinula clathrata
Lamarck, is in the Muséum d’Histoire Naturelle,
Geneva, no. 1101/14/1 (PI. 28, fig. 1). According
to Rosalie de Lamarck’s marginal annotations in
her father’s copy of the “Histoire naturelle des
animaux sans vertebres,” only a single specimen
was present in Lamarck’s collection at the time
of description. In 1822 Lamarck gave the size of
his specimen as 13% lignes [= 30.4mm], and this
dimension agrees with the larger specimen,
which is considered the holotype, but not the
smaller 29.7mm specimen which accompanies it.
This latter specimen has probably been added at
a later date. The type locality here designated is
Tuamotu Islands (after Reeve, 1846), specifically
Raroia Island (from which there are specimens in
the ANSP).
Nomenclature—This species seems to have
been confused with Drupa rubuscaesius Roding,
and D. rubusidaeus Réding, by modern authors.
Hedley (1913) initiated the confusion by suggest-
ing that Ricinula clathrata Lamarck, R. speciosa
Dunker and Purpura spathulifera Blainville, were
synonyms of Drupa rubuscaesius Roding. Kira
(1954, pl. 23, fig. 9; 1962, pl. 24, fig. 9) illus-
trates a specimen of Drupa clathrata with an im-
mature lip and calls it in the first instance
D. rubuscaesium Roding, and in the second in-
stance D. rubsidaeus (sic) Roding. Kaicher (1957,
[14-846]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13
Drupa_ 33
Plate 29. Geographical distribution of Drupa (Ricinella)
clathrata clathrata (Lamarck) and its subspecies D. (R.)
clathrata miticula (Lamarck). Open circles are literature
records.
pl. 3, fig. 15) cites D. clathrata as a synonym of
D. rubuscaesia Roding, although it is not clear
to which species her figures refers. As discussed
earlier, Drupa rubuscaesius Roding, is a synonym
of D. ricinus (Linnaeus), for Réding refers to
Martini’s figure of the yellow apertured form of
that species.
Records—PHILIPPINES: Gigmoto, Catanduanes Id.
(AMNH; DMNH; AIM); Borongan village, E. Samar Id.
(ANSP); Marinduque Id. (USNM). FORMOSA (Taiwan):
Chaiting (Janowsky coll.). RYUKYU ISLANDS: Yakushima:
Onna Beach, N. of Naha, Okinawa; Bolo Point, Zampa Mis-
aki, Okinawa (all USNM). JAPAN: Hackijo Id., 275 mi S.
of Tokyo; Tosa, Shikoku; Kagoshima, Kyushu (all ANSP);
Oshima, Osumi (USNM). BONIN ISLANDS: Chichi Shima
(ANSP); Ani Jima (USNM). MARIANAS: Lagunan Tanapag,
Saipan Id.; Ngargersul Id. (both ANSP); off Leprosarium,
SW Tinian Id. (MCZ); Aspurguan, Guam Id.; Ypao Point,
Maug Id. (USNM). WAKE ISLAND: (AMNH). MARSHALL
ISLANDS: (many atolls, see map); Eniwetok: Bikini; Ronge-
lap; Kwajalein; Jaluit (all USNM). SOLOMON ISLANDS:
Choiseul Bay, Choiseul Id. (AMNH). NEW HEBRIDES:
Pango Point, Efate Id., (AIM). LOYALTY ISLANDS: Lifu
(USNM; AMS). NEW CALEDONIA: Touho (AMNH). GIL-
BERT ISLANDS: Kingsmill Id. (USNM). FIJI ISLANDS:
Cuvu Beach, S. Viti Levu (AMNH); Rat Tail Passage, Suva
reef, S. Viti Levu (WOC coll.); SE of Onea Driki, Lau group
(USNM). TONGA ISLANDS: Ha’ateiho reef, Tongatapu;
Niuafo’ou (both USNM). NIUE ISLAND: (DMNH); Oneone
reef; Utuko reef (both AMNH): Alofi (USNM). SAMOA IS-
LANDS: Tutuila Id. (ANSP); Swan’s Id. (MCZ). Lalomalava,
Savaii (DMNH). COOK ISLANDS: Bird’s Id., Palmerston
Atoll; North Id., Palmerston Atoll (both USNM); Avaavaroa
Passage, S. Rarotonga (ANSP). SOCIETY ISLANDS:
Moorea; Venus Point, Tahiti (both AMNH); Punaavia, Tahiti
(ANSP). TUAMOTU ISLANDS: Tacume: Raroia (both
ANSP); Anaa Id. (AMS); Makemo: Tikahau; Vahitahi; Maka-
tea (all USNM). MARQUESAS ISLANDS: Atuona Bay, Hiva-
Oa Id. (ANSP). PITCAIRN ISLAND: (AMNH). LINE IS-
LANDS: Caroline Id. (ANSP); Jarvis (DMNH).
Drupa clathrata miticula (Lamarck, 1822)
(Pl. 2, figs. 19, 20; Pls. 28, 30)
Range—Madagascar to the Island of Timor in
Indonesia.
Remarks—First described by Lamarck in 1822,
the identity of this Indian Ocean form appears to
have escaped notice for over a hundred years
and in that interval has been recorded as typical
Drupa clathrata. It lacks the brown coloration of
the columella and inner and outer lips that char-
acterize the nominate subspecies. D. miticula is
also a smaller shell, with shorter spines. The In-
dian Ocean populations have a more pronounced
purple color in the aperture than those of the
nominate subspecies which is lavender to whitish
purple. The exterior lacks the black spines of
D. morum.
Habitat—In tide pools, basalt rock, weed and
some coral, and on sand and grass, from 0 to 8
feet. In the Cocos-Keeling Islands the species
was found in “strong surf on a large boulder on
the northern seaward reef at Horsburgh Island”
(V. Orr Maes, personal communication). On
Christmas Island it was found among Caulerpa
I 0.1 mm
Plate 30. Radula of Drupa (Ricinella) clathrata miticula
(Lamarch). Half a transverse row; Greta Beach, Christmas
Id., Indian Ocean.
[14-847]
34. Ricinella
Emerson and Cernohorsky
mats and in rock and tide-pools (leg. A. Slack-
Smith and A. Patterson).
Description—Shell 16 to 38 mm (% to 12 inches)
in length, ovate, globose, spire short, acuminate.
Body whorl large with six transverse rows of
short tubercles. Interstices between tubercles
striated with four to six rows of closely spaced
granules. Aperture moderately wide, oval, over
three-quarter of shell length. Outer lip crenulated
between tubercles, inner margin dentate with six
small white teeth from which six conspicuous
white raised lines run into the aperture. Inner lip
enameled with three to four plait-like ridges pro-
jecting into the aperture. Columella excavated
posteriorly. Posterior siphonal canal open,
obliquely curved toward the apex. Color pale
brown on the exterior, interior of aperture
purple.
In the radula examined, one of the side-cusps
of the rachidian was trifid while the other one
was simple; there were 2 lateral denticles apart
from the end-cusps.
Measurements (mm)—(including spines; all
specimens with a mature lip).
length width
38.0 31.3 Christmas Id., Indian Ocean
34.3 28.0 S. Mahébourg, Mauritius
32.5 26.0 S. Mahébourg, Mauritius
26.3 21.0 Lectotype of miticula Lamarck
17.0 13.8 Christmas Id., Indian Ocean
16.5 14.5 Souillac, Mauritius
Muricidae
Synonymy—
1822 Ricinula miticula Lamarck, Hist. Nat. anim. s. verte-
bres, vol. 7, p. 231 (no locality given); 1832 Blain-
ville, Nouv. Ann. Mus. d’Hist. Nat. Paris, ser. 3, vol. 1,
p. 211; 1844 Deshayes & Milne-Edwards, Hist. nat.
anim. s. vertebres, ed. 2, vol. 10, p. 48 (refers to
Kiener, fig. 5, junior = fig. 5a, 5a).
1835 Purpura clathrata Lamarck, Kiener (pars), Spéc. gén.
icon. coq. viv., vol. 8, p. 15, pl. 3, figs. 5a, 5a (non
Ricinula clathrata Lamarck, 1816).
1968 Drupa clathrata Lamarck, Taylor, Phil. Trans. Roy. Soc.
London, ser. B, vol. 254, p. 201 (non Ricinula clath-
rata Lamarck, 1816).
Types—Two syntypes of Ricinula miticula are in
the Muséum d Histoire Naturelle, Geneva. The
slightly larger specimen, length 26.3 mm _ (PI.
28, fig. 3), no. 1101/13/1, is here selected as the
lectotype of R. miticula. No type locality was
given, and Mahébourg, Mauritius, is here desig-
nated as the type locality (specimens in AMNH).
Nomenclature—Kiener (1835) regarded _ Rici-
nula miticula of Lamarck to be a juvenile of
R. clathrata, but his figures are typical R. clath-
rata miticula.
Records—(Specimens): MAURITIUS: Gris Gris, 1 mi. ESE
of Souillac; Vacoas Point, 3 mi. S. of Mahébourg; Point
Pimente, N. side Arsenal Bay; Pointe Fayette; Caves Point
(all ANSP); Mahébourg (AMNH); near Port Louis (MCZ).
COCOS-KEELING ISLANDS: N. end of Horsburgh Id
(ANSP). CHRISTMAS ISLAND: Lily Beach; Greta Beach;
Dolly Beach (all WAM). INDONESIA: Timor (AMS).
Records—(Literature—identified as “clathrata”): SEY-
CHELLES ISLANDS: Coetivy Id. (Melvill, 1909, p. 104);
Mahé (Taylor, 1968, p. 201). MADAGASCAR: (Dautzenberg,
1923, p. 38). MAURITIUS: (Viader, 1937, p. 32). REUNION
ISLAND: (Deshayes, 1863, p. 115).
[14-848]
May 30, 1973
Subgenus Drupina Dall, 1923
Type: Drupa grossularia Réding, 1798
Shell sub-ovate, strong, heavy, flattened dorso-
ventrally, spire short; whorls ribbed spirally with
inconspicuous nodules, surface sculptured with
minute imbricated scales; columella doubly _pli-
cated axially, outer lip with two well developed
marginal processes. Operculum typical for genus.
Radula with a broad but low rachidian which
has from 13-18 cusps; the central cusp may be
large or small and the 2 flanking side-cusps are
tridentate and usually smaller than the central
cusp. The lateral teeth are small and_ slender,
with a smaller and more rounded base than in
Drupa sensu stricto.
The 2 members of this subgenus have a por-
tion of the outer lip expanded as two conspic-
uous lobate processes. The process develops as
the individual reaches maturity and thus is not
laid down and re-absorbed by the mantle as the
shell grows; instead the processes are thickened,
extended and often bifurcated. Both species of
the subgenus Drupina are Indo-Pacific in dis-
tribution.
Synonymy—
1923 Drupina Dall, Proceedings of the Academy of Natural
Sciences of Philadelphia, vol. 75, p. 303. Type-species
by original designation: Ricinula digitata Lamarck,
1816 [= Drupa grossularia Ré6ding, 1798).
| Ochi
Plate 31. Radula of Drupa (Drupina) grossularia Roding.
Half a transverse row; Fiji Islands.
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 35
Drupa grossularia Réding, 1798
(Pl. 2, figs. 23, 24; Pls. 31, 32)
Range—From the Cocos-Keeling Islands in the
Indian Ocean to West Australia and throughout
the Pacific to Hawaii and the Marquesas Islands.
Remarks—This is a very distinctive species with
the large digitate processes and solid yellow
aperture distinguishing it from all other mem-
bers of the genus except Drupa lobata (Blain-
ville), a dark brown apertured form inhabiting
the Indian Ocean.
Habitat— Lives clinging to rocks exposed at low
tide or ina few feet of water on windward rather
than leeward reefs. Demond (1957) records a
specimen taken alive at a depth of 32-38 feet
in a lagoon west of Saipan.
Description—Shell 18 to 33 mm (% to 14 inches)
in length, ovate, spire very short, body whorl
large. Whorls spirally ribbed with low rounded
nodules most common; nodules often as siphon-
Plate 32. Drupa (Drupina) grossularia Réding.
Fig. 1. Lectotype figure of Drupa grossularia Réding, from
Martini, 1777, Syst. Conchylien-Cabinet, vol. 3, pl. 102,
fig. 978.
Fig. 2. Type figure of Purpura laurentiana Petit de la
Saussaye, from Journal de Conchyliologie, 1850, vol. 1,
pl. 13, fig. 2; Pacific Ocean (20.0 x 16.0 mm-—yjuvenile
specimen).
Fig. 3. Lectotype of Ricinula digitata Lamarck (MHNCG no.
1101/16/1; 22.4 x 22.0 mm)
[14-859]
36 = Drupina
ous tubercles near the margin of the aperture.
Sculpture of crisp, well-defined scales; detail of
sculpture retained only in well preserved indi-
viduals. Aperture in adult stage, linear, con-
stricted by 5 or 6 singularly arranged, close-set
teeth projecting from the outer lip, and a col-
umellar callus with from 2-5 inconspicuous pli-
cations. Columella doubly folded axially. Aper-
ture of juvenile specimens expanded. In adults,
siphonous, digitate processes extend from the
first and second ribs of the body whorl. The proc-
esses tend to bifurcate at maturity; canal of the
upper process canaliculate, lower canal generally
sealed in the adult stage. Exterior of shell white
or cream, aperture yellow to orange, denticles of
outer lip white, columellar plications faintly whit-
ish. Operculum typical for the genus, dark
orange-brown in color.
The radula is of a considerably modified dru-
pine type and has been described in the sub-
generic diagnosis. The size and length of the cen-
tral tooth and accessory lateral denticles are
quite variable.
Measurements (mm)—(including digitations; all
specimens with a mature lip).
length width
32.7 33.6 Limu, Niue Island
32.0 30.9 S. Luzon Id., Philippines
23.2 22.2 Kavieng, New Ireland
22.4 22.0 Lectotype of digitata Lamarck
18.7 18.3 Bougainville, Solomon Ids.
Synonymy—
1685 Lister, Hist. Syn. Meth. Conchyliorum, pl. 804,
fig. 12.
1758 Seba, Locupl. rer. nat. thes. descriptio, vol. 3, pl.
60, fig. 48.
mon «
binomial).
1791 Murex neritoideus Gmelin (pars), Systema Naturae,
ed. 13, p. 3537 (refers to Seba, op. cit., Lister, op. cit.,
and Martini, figs. 978, 979 only) [non Linnaeus,
1767].
1798 Drupa grossularia Réding, Museum Boltenianum, p. 55
(refers to Martini, op. cit.) [no locality given]; 1913
Hedley, Nautilus, vol. 27, no. 7, p. 80; 1957 Kaicher,
Indo Pacific Sea Shells (Muricacea, Buccinacea), pl. 4,
fig. 4; 1970 Salvat, Cahiers du Pacifique, no. 14,
p. 46.
1816 Ricinula digitata Lamarck, Tableau Encyclopédique
Méthodique, p. 2, pl. 395, figs. 7a, b (no locality
given); 1822 Lamarck, Hist. nat. anim. s. vertébres,
vol. 7, p. 232; 1827 Crouch, Ilust. Introd. Lamarck’s
Conchology, p. 36, pl. 18, fig. 8; 1842 Reeve (pars),
Conchologia Systematica, vol. 2, p. 215, pl. 256,
fig. 3 only; 1846 Reeve, Conchologia Iconica, vol. 3,
pl. 1, fig. 2a (Lord Hood Id. = S. Marutea Id.); 1859
Chenu, Manuel de Conchyliologie, vol. 1, p. 168,
text fig. 815; 1880 Tryon, Manual of Conchology,
vol. 2, p. 185, pl. 56, fig. 191 and pl. 57, fig. 203;
1933 Dautzenberg & Bouge, Journal de Conchyliol-
ogie, vol. 77, p. 237.
Emerson and Cernohorsky
Muricidae
1842 Purpura monstruosa Lesson, Rev. Zool Cuvierienne,
vol. 5, App. p. 103 [Gambier Islands].
1823 Murex fimbriatus Mawe. Linn. Syst. Conchology,
p. 131, pl. 26, fig. 4 (non Brocchi, 1814; nec
Lamarck, 1822).
1825 Murex ricinus Wood, Index Testaceologicus, pl. 26, fig.
5Sla (non Linnaeus, 1758).
1832 Purpura digitata Lamarck, Blainville, Nouv. Ann. Mus.
(Hist. Nat. Paris, ser. 3, vol. 1, p. 210; 1833 Quoy
& Gaimard, Voyage L’Astrolabe, vol. 2, p. 578, pl. 39,
figs. 20-22 (shell, animal and operculum) [Carteret
Harbour, New Ireland]; 1835 Kiener, Spéc. gén. icon.
coq. viv., vol. 8, p. 16, pl. 3, figs. 6, 6a.
1850 Purpura laurentiana Petit de la Saussaye, Journal de
Conchyliologie, vol. 1, no. 4, p. 403, pl. 13, fig. 2
(Pacific Ocean) [juvenile specimen].
1853 Pentadactylus grossularius Bolten, H. & A. Adams,
Gen. Rec. Mollusca, vol. 1, p. 129 and vol. 3, pl. 13,
fig. 6c; 1875 Troschel, Gebiss d. Schnecken, vol. 2,
p. 133, pl. 13, fig. 1 (radula).
1880 Ricinula hystrix var. laurentiana Petit, Tryon, Manual
of Conchology, vol. 2, p. 184, pl. 56, fig. 192 (juvenile
specimen).
1908 Pentadactylus (Pentadactylus) digitatus Lamarck, Horst
& Schepman, Cat. Syst. Moll. Mus. Hist. Nat. Pays-
Bas, vol. 13, p. 157.
1929 Drupina grossularia Rodding, Iredale, Mem. Queens-
land Museum, vol. 9, pt. 3, p. 290; 1961 Rippingale
& McMichael, Queens]. & Gt. Barrier reef Shells, p.
102, pl. 13, fig. 2; 1962 Kira, Shells west. Pacific in
color, p. 62, pl. 24, fig. 3; 1965 Arakawa, Venus:
Jap. Journ. Malacology, vol. 24, no. 2, p. 116, pl. 13,
figs. 8-10 (radula); 1969 Cernohorsky, Veliger, vol.
11, no. 4, p. 303, pl. 48, fig. 11 (shell), text fig.7
(radula); 1971 Wilson & Gillett, Australian Shells,
p. 92, pl. 61, figs. 5, 5a; 1971 Kay, Pacific Science,
vol. 25, pp. 263, 275.
1952 Sistrum digitatum Lamarck, Morris, Field Guide to
shells Pacific coast and Hawaii, p. 187, col. pl. 5,
fig. 5; pl. 39, fig. 3.
1965 Drupina glossularia (sic) (R6ding), Wu, Bull. Inst. Zool.
Acad. Sinica, vol. 4, p. 99, text figs. 20, 21 (radula);
1967 Habe & Kosuge, Stand. Book Jap. shells in
color, vol. 3, p. 70, pl. 27, fig. 22 (invalid emenda-
tion).
1968 Drupa (Drupina) grossularia Réding, Orr Maes, Pro-
ceedings of the Academy of Natural Sciences of Phil-
adelphia, vol. 119, no. 4, p. 130.
Types—The holotype of Drupa grossularia is
no longer traceable and the specimen figured by
Martini on plate 102, figs. 978, 979 (Pl. 32, fig.
1) which was cited by Réding, is here designated
as the lectotype of the species. Two syntypes of
Ricinula digitata Lamarck, are in the Muséum
dHistoire Naturelle, Geneva, and the 22.4mm
long specimen, no. 1101/16/1, which most
closely corresponds to Lamarck’s cited dimen-
sions, is here selected as the lectotype (PI. 32,
fig. 3). The holotype of Purpura laurentiana Petit
de la Saussaye, is in the Muséum National d’His-
toire Naturelle, Paris (Journ. de Conchyliologie
coll.). No type locality was given by Réding for
D. grossularia, and the earliest record of Car-
teret Harbour, New Ireland, by Quoy & Gaimard,
[14-860]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13
Drupa 37
160°
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2 a 60" BO" -_ roe 20 y40" 160" : 180 Teo" 140 120" Too? 3°
Plate 33. Geographical distribution of the species Drupa
(Drupina) grossularia Réding, in the Pacific and East In-
dian Oceans (full circles) and D. (D.) lobata (Blainville)
in the Indian Ocean (half-open circles).
1833, is here designated as the type locality, as
well as for Purpura digitata Lamarck. Two syn-
types of Purpura monstruosa Lesson, which are
said to be “typical” examples of D. grossularia,
are in the Muséum National d’Histoire Naturelle,
Paris (teste G. Richard).
Nomenclature—There can be no doubt as to
Réding’s concept of Drupa grossularia, as he,
Schumacher and Lamarck, all cite identical figures
in Martini, 1777. Petit’s Purpura laurentiana is a
young specimen in which the mature lip has not
started to form and has the whorls covered with
a foreign growth (PI. 32, fig. 2). A similar juve-
nile is illustrated on PI. 2, fig. 24.
Records—COCOS-KEELING ISLANDS: S. end of Home Id.
(ANSP). INDONESIA: Marudu Bay, N. Borneo (ANSP)
Malawali Channel, N. Borneo (AMNH; USNM); Bali; Mor-
otai Id. (both AMS); Oong Bay, Mandidi Id., Moluccas
(MCZ); Pulau Pombo, Wasi, Ambon Id. (FMNH). PHIL-
IPPINES: Recorded from many localities throughout the
Archipelago: Luzon; Mindoro; Catanduanes; Busanga; Bala-
bac; Sulu Archipelago (AMNH; ANSP; MCZ; USNM); Boron-
gan village, E. Samar Id. (AIM). FORMOSA (Taiwan): Now
Wow (USNM). RYUKYU ISLANDS: Sobe; Kadena; Mekan-
iko; Onna reefs; Bolo Point, all Okinawa (all AMNH);
Miyako (FMNH). JAPAN: Hachijo Id., 275 mi. S. of Tokyo;
Tosa, Shikoku; Osima, Osumi (all ANSP). MARIANAS:
Lagunan Tanapag, Saipan; Agat Bay, Guam (both ANSP);
Tinian Id. (MCZ). PALAU ISLANDS: Helen reef, Koror;
Babelthuap (both ANSP). CAROLINE ISLANDS: S. of Gar-
abayo; Kayangel; Rattakadokoru; Ngaruangl; Ngangersul;
Ponape; Yap (all ANSP); Elato Atoll; Lamotrek Atoll; Sata-
wal Atoll; Kapingamarangi (all USNM); Lukunor (AMNH).
MARSHALL ISLANDS: Eniwetok Atoll; Bikini Atoll; Kwaja-
lein Atoll (all USNM); Arno Atoll (AMNH). ADMIRALTY
ISLANDS: Manus Id. (DM). NEW BRITAIN: Rabaul
(AMNH,; USNM). NEW IRELAND: Kavieng (AMNH). NEW
GUINEA: Aoeri Ids., Geelvink Bay; SW Biak, Schouten Ids.;
Wooi Bay, Japen Id.; 1 mi. NE Mioes Woendi, Padaido Ids.
(all ANSP); Samarai, Papua; Port Moresby, Papua (both
AMNH); Milne Bay, Papua (USNM). AUSTRALIA: Torres
Strait (Shirley, 1912, p. 102); Queensland: Michaelmas Cay,
off Cairns; Bramble Bay, off Lucinda (both AMS); West
Australia: Barrow Id. (Wilson & Gillet, 1971, p. 92. SOLO-
MON ISLANDS: Vanikoro Id., Santa Cruz group; Reef Id.,
Santa Cruz group (both AMS); Kieta, Bougainville Id.; Lutee,
Choiseul Id.; Ataa, N. Malaita Id. (all AMNH); Ugi Id.,
Shortland group (USNM); Ticopia Id., (AIM); NEW HEB-
RIDES: Meli Id., SW Efate Id.; Pango Point, Efate Id. (both
AIM); Pentecost Id. (Powell coll.). LOYALTY ISLANDS: Lifu
(AMS). NEW CALEDONIA: Touho (AMNH); Bourail; 7 mi.
SW Gatope Id., Voh (both ANSP). FIJI ISLANDS: 3 mi.
NE Tunuloa, Vanua Levu (MCZ); Suva Harbour, S. Viti
Levu; Ogea Levu, Ogea, Lau group (both USNM); Wadigi
Id., Mamanuca group (WOC coll.). GILBERT ISLANDS:
Abaiang (MCZ); Onotoa Atoll, Kingsmill Ids. (USNM). EL-
LICE ISLANDS: Funafuti (AMS; AIM). WALLIS & FUTUNA:
Nukuhifala, Wallis Id.; W. coast of Uvea, Wallis Ids.; E. side
of Faioa, Wallis Ids. (all USNM); Anse de Sigave, Hoorn
Id., Futuna Ids. (USNM). SAMOA ISLANDS: Asau Harbour,
Savaii (USNM):; Vailele Bay, Upolu Id. (ANSP); Satalo Id.,
Upolu Id. (AIM); Apia, Upolu Id. (Powell coll.); Pago Pago,
Tutuila Id. (AMNH; MCZ). TONGA ISLANDS: Ha’ateiho,
Tongatapu (USNM). NIUE ISLAND: Limu (AMNH); Alofi
(USNM). PHOENIX ISLANDS: Enderbury Id. (USNM).
COOK ISLANDS: Akamaru, Manihiki Atoll (ANSP); Bird’s
Id. and Tom’s Id., Palmerston Atoll; Motu Akaiami, Aitutaki
(all USNM); Mauke (ANSP); several localities on Rarotonga
(MCZ; USNM; AIM). AUSTRAL ISLANDS: Rurutu; Raivavae
(both USNM). SOCIETY ISLANDS: N. of Fare, Huahine;
Faredine, NW Moorea; Papeete, Tahiti (all USNM). TUA-
MOTU ISLANDS: Raroia Id.; Lord Hood Id. [= S$. Marutea
Id.] (both AMNH); Makemo Id.; Toau Atoll (both ANSP);
Nengonengo Id. (USNM). GAMBIER ISLANDS: Mangareva
Id. (USNM). MARQUESA ISLANDS: (USNM; ANSP). LINE
ISLANDS: Palmyra Id.; Flint Id.; Christmas Id. (all ANSP);
Fanning Id.; Kingman reef; Washington Id.; Jarvis Id. (all
Kay, 1971, p. 275). HAWAIIAN ISLANDS: Kure Id.; Midway
Id. (both USNM; Makua, Oahu (Adams, 1967, p. 4).
Cm nai
L O.1 am J
Plate 34. Radula of Drupa (Drupina) lobata (Blainville). Half
a transverse row; Veeckens Bay, S. Pagi Id., Indonesia.
[14-861]
38 = Drupina
Drupa lobata (Blainville, 1832)
(PI. 2, figs. 25, 26; Pls. 34, 35)
Range—From the Red Sea and East Africa
through the Indian Ocean to Thailand, Sumatra
and West Australia.
Remarks—Despite its closeness to Drupa gross-
ularia, the present chocolate-mouthed species
seems to be a distinct form which replaces the
yellow-mouthed species in the Red Sea and In-
dian Ocean. Their ranges are known to overlap
in the Cocos-Keeling Islands where the two
forms were collected at Home Island and no
intermediates were found (Ostheimer and Orr
Maes leg.), and along the coast of West Australia
(specimens in West Australian Museum).
Habitat—On intertidal wave-swept reefs. In the
Cocos-Keeling Islands it was found on the tops
and sides of rocks near shore on the seaward
reefs, and on the seaward ends of passes (Orr
Maes, personal communication).
Description—Shell 18 to 32 mm (% to 1% inches)
in length, very similar to Drupa grossularia Rod-
ing, but differing in having a chocolate-brown
aperture, a brownish dorsum, and in having a
wider, more lobate digitate process extending
from the first rib on the body whorl. This proc-
ess shows no tendency to bifurcate as it does in
D. grossularia, and the canal remains open in
the adult.
The radula is similar to that of D. grossularia;
the side-cusps of the rachidian are short, broad
and trifid, and are followed by 6 moderately
deeply rooted lateral denticles.
Measurements (mm)—(including digitations; all
specimens with a mature lip)
length width
33.2 33.0 Mauritius
28.6 27.8 Kiwengwa, Zanzibar
26.5 26.3 Syntype of dactyloides Schumacher
26.2 27.5 Eilat, Gulf of Aqaba
18.5 17.5 Zanzibar
Synonymy—
1817 Ricinella dactyloides Schumacher, Essai nouv. systéme,
p. 241 (refers to Martini, 1777, vol. 3, pl. 102, figs.
978, 979 = Drupa grossularia Riding) [no locality
given] (nomen oblitum).
1823 Ricinula digitata Lamarck, Sowerby, Gen. Rec. foss.
shells, pt. 18, pl. 235, figs. 3, 4 (no locality given)
[non Lamarck, 1816].
Emerson and Cernohorsky
Muricidae
1832 Purpura lobata Blainville, Nouv. Ann. Mus. d’Hist.
Nat. Paris, ser. 3, vol. 1, p. 210, pl. 9, fig. 7 (no
locality given); 1835 Kiener, Spéc. gén. icon. coq. viv.,
vol. 8, p. 18, pl. 3, fig. 7.
1842 Ricinula digitata var. Lamarck, Reeve (pars), Conchol-
ogia Systematica, vol. 2, p. 215, pl. 256, fig. 4 only
(non Lamarck, 1816).
1844 Ricinula digitata var. fusca “Sowerby,” Deshayes &
Milne-Edwards, Hist. Nat. anim. s. vertebres, ed. 2,
vol. 10, p. 53 (no locality given) [published in syn-
onymy of R. lobata Blainville—refers to Sowerby,
1823, pl. 235, fig. 4] (non R. fusca Kiister, 1862).
1846 Ricinula digitata var. B. Reeve, Conchologia Iconica,
vol. 3, pl. 1, fig. 2b (Seychelles Ids.) [non Lamarck,
1816].
1880 Ricinula digitata var. lobata Blainville, Tryon, Manual
Conchology, vol. 2, p. 185, pl. 57, fig. 205.
1896 Ricinula lobatus Blainville, Shopland, Journ. Bombay
Soc. Nat. Hist., vol. 10, p. 220.
1903 Sistrum digitatum (var. lobata) E. A. Smith in Gardi-
ner, Fauna & Geog. Maldive & Laccadive Archi-
pelago, p. 609.
1919 Drupa digitata var. lobata Blainville, Cooke, Proc.
Malac. Soc. London, vol. 13, pt. 4, p. 101 (descrip-
tion of radula); 1937 Viader, Mauritius Inst. Bull.
vol. 1, pt. 2, p. 32.
1913 Ricinula lobata Blainville, Hedley, Nautilus, vol. 27.
no. 7, pp. 79, 80; 1922 Hornell, Madras Fish. Dept.
Bull., no. 6, p. 217.
1950 Drupa (Drupina) grossularia lobata Blainville, Abbott,
Bull. Raffles Museum, vol. 22, p. 80.
1956 Drupa (Drupina) lobata (Blainville), Franc, Ann. L’'Inst.
Océanog. Monaco, N.S. 32, p. 37 (Ile Abulat, Red
Sea); 1967 Orr Maes, Proc. Acad. Nat. Sci. Philadel-
phia, vol. 119, no. 4, p. 130, pl. 11, fig. E.
1961 Drupa lobata Spry, Tanganyika Soc. Notes & Record,
no. 56, p. 21, pl. 7, fig. 142.
1969 Drupina lobata (Blainville), Cernohorsky, Veliger, vol.
11, no. 4, p. 303 1970 Heinicke, Hawaiian Shell
News, vol. 18, no. 7; p. 6, text fig.; 1971 Wilson &
Gillett, Australian Shells, p. 92, pl. 61, fig. 4 (Pt.
Cloates, West Australia).
Types—The type specimen of Purpura lobata
Blainville, is presumably in the Muséum National
dHistoire Naturelle, Paris. Five probable syn-
types of Ricinella dactyloides Schumacher, are
in the Zoological Museum, Copenhagen. Four of
these specimens have the letters “Sp” marked
either in the aperture or on the dorsum, and these
originated from the Spengler collection. One
specimen is marked “Sch” [= Schumacher] in-
side the aperture (PI. 35, fig. 1). The type: local-
ity of D. lobata is here designated as Mogadiscio,
Somalia.
Nomenclature—When Schumacher described
Ricinella dactyloides, his diagnosis consisted of
only the three words “labio externo digitato”;
for an illustration he referred to Martini’s figures
978, 979, which represent Drupa_ grossularia
Réding. However, the extant and probable syn-
types of Ricinella dactyloides are referable to the
species Drupa lobata (Blainville), and Schu-
macher’s taxon would in effect have 15 years
[14-862]
May 30, 1973
Plate 35. Drupa (Drupina) lobata (Blainville).
Fig. 1. Probable syntype of Ricinella dactyloides Schu-
macher; marked;‘Sch[umacher]” inside aperture (ZMC;
26.5 x 26.3 mm).
Fig. 2. Probable syntype of R. dactyloides Schumacher;
marked#‘Sp[engler]” inside aperture (ZMC; 26.8 x 25.8
mim).
priority over Blainville’s. Since Schumacher’s
name has not once been applied to a taxon as
the valid name during the last 50 years, it is con-
sidered to be an unused senior synonym. The
taxon Purpura lobata Blainville, however, has
been in general current use during the preceding
INDO-PACIFIC MOLLUSCA, vol. 3, no. 13 Drupa 39
fifty years, and has been used by 5 different
authors in 10 publications (see Declaration 43 of
the ICZN; Bull. Zool. Nomencl., vol. 27, pts.
3/4, p. 135).
Hedley (1913) suggested that the name fusca
Deshayes & Milne-Edwards, 1844, be applied to
the present species on thé mistaken belief that
Blainville had proposed Purpura lobata for the
yellow-apertured Drupa digitata (Lamarck) [=
D. grossularia Roding]. Blainville (1832, p. 210)
clearly states: “couleur d'un’ brun-marron en
dehors et & la circonférence de louverture,
blanche en dedans”; the type-figure given by
Blainville (1832, pl. 9, fig. 7) is also an excellent
representation of the dark brown apertured form,
despite the lack of cited locality.
Records—RED SEA: Eilat, Gulf of Aqaba, Israel(A. Hadar;
K. Haim; AMNH; DMNH); He Abulat (Franc. 1956, p. 37);
Jidda, Saudi Arabia (DMNH) GULF OF ADEN: Aden (Shop-
land, 1896, p. 220). EAST AFRICA: 9 mi. N. of Mogadiscio,
Somalia (ANSP); at 19 km marker, N. of Mogadiscio, So-
malia (AMNH); Diani Beach, Kenya (Heinicke, 1970, p. 7);
15 mi. SSE of Dar-es-Salaam, Tanzania; 4 mi. ESE of Dar-
es-Salaam, Tanzania (both MCZ); Mozambique City, Mozam-
bique (ANSP). ZANZIBAR: Pange Id. Kiwengwa;
Mangapivani (all ANSP). SEYCHELLES: Beau Vallon Beach,
Mahé (ANSP). MADAGASCAR: Grande Recife, W. end of
ship pier, Tuléar; Grande Recife, W. of airport, Tulear (both
MCZ). REUNION: (Deshayes, 1863, p. 115). MAURITIUS:
NW side of Tamarin Bay (ANSP). MALDIVE ISLANDS:
Imma Id., N. Male Atoll; Fodiffolu Atoll; Ari Atoll (all
ANSP). LACCADIVE ISLANDS: (Hornell, 1922, p. 217).
THAILAND: Goh Phi Phi; Goh Huyong, Similan Ids. (both
USNM). COCOS-KEELING ISLANDS: N. tip West Id.; S.
end of Direction Id.; S$. end Home Id. (all ANSP). CHRIST-
MAS ISLAND: (Tomlin, 1935, p. 79). INDONESIA: Pulau
Siburu, N. of Sipora, S.W. Sumatra;Pulau Bai, Batu group,
off Sumatra; W. shore Veeckens Bay, S. Pagi Id. (all USNM).
WEST AUSTRALIA: W. of Ningaloo homestead, Pt. Cloates,
22°42’S and 113°39’E (WAM).
[14-863]
40 Index
Emerson and Cernohorsky
INDEX TO DRUPA NAMES IN VOL. 3 NO. 13
The number following the name refers to the
pagination found at the top of the page. The col-
umn at right is the looseleaf pagination. Al
new names are in bold face type.
aesculus Réding, 9
albolabris Blainville, 20
album Montfort, 20
arachnoides Lamarck, 20
Azumamorula, 8
baylei Coss. & Lambert, 5
bollonsi Suter, 7
botroides Réding, 9
Canrena Link, 14
chamaemorus Réding, 9
Condonia Hertlein, 8
cornus Roding, 9
clathrata Lamarck, 31
dactyloides Schumacher, 38
digitata Lamarck, 36
Drupa Réding, 14
Drupina Dall, 35
Drupinae, 13
elegans Broderip & Sowerby, 24
fimbriatus Mawe, 36
fragun, Roding, 28
fusca Deshayes & Edwards, 38
glans Réding, 9
globosa Morch, 16
globosum Martini, 16
grossularia Réding, 35
hadari Emerson & Cernohorsky, 23
hannai Howe, 8
hericinus Martini, 28
hippocastanum Wood, 28
horrida Lamarck, 16
hystrix Linnaeus, 20
hystrix auctt., 28
iodostoma Lesson, 18
jodostoma Boettger, 19
14-809
14-824
14-824
14-824
14-808
14-805
14-807
14-809
14-818
14-809
14-808
14-809
14-845
14-862
14-860
14-818
14-859
14-817
14-828
14-860
14-842
14-862
14-809
14-820
14-820
14-859
14-827
14-808
14-842
14-842
14-820
14-824
14-842
14-822
14-823
laurentiana Petit de la Saussaye, 36
lobata Blainville, 38
mancinella Roding, 9
miticula Lamarck, 33
monstrosa Lesson, 37
morum Roding, 15
muricina Réding, 9
mutica Lamarck, 8
neritoidea Link, 16
neritoideus Linnaeus, 16
neritoideus Mawe, 20
nodosa Linnaeus, 16
nodus Gmelin, 28
Pentadactylus Morch, 14
pisolina Lamarck, 8
purpurata Schumacher, 28
Purpurinae, 13
puruensis K. Martin, 5
reeveana Crosse, 28
rhombiformis K. Martin, 5
Ricinella Schumacher, 27
Ricinula Lamarck, 14
Ricinulus Demarest, 14
ricinus Linnaeus, 19
rubuscaesius Réding, 20
rubuscestus Dall, 20
rubusidaeus Roding, 27
Sistrum Montfort, 14
smithi Brazier, 8
spathulifera Blainville, 28
speciosa Dunker, 30
styriaca Stur, 5
Thaidinae, 13
trapa Réding, 9
tribulus Roding, 20
uva Réding, 9
violacea Schumacher, 16
vitiensis Pilsbry in P. & B., 9
walkerae Pilsbry & Bryan, 9
[14-864]
Muricidae
14-860
14-862
14-809
14-847
14-861
14-819
14-809
14-808
14-820
14-820
14-824
14-820
14-842
14-818
14-808
14-842
14-817
14-805
14-842
14-805
14-841
14-818
14-818
14-823
14-824
14-824
14-841
14-818
14-808
14-842
14-844
14-805
14-817
14-809
14-824
14-809
14-820
14-809
14-809
INDO-PACIFIC WILLIAM H. DALLE
MOLLUSCA —oSsiii'SEEs
volume 3, no. 14
THIS SECTION CONTAINS
1. The Genus Gabrielona (Phasianellidae) in the Indo-Pacific and West Indies.
By Robert Robertson. (Put in Binder 1, just after the guide tab “Pha-
sianellidae”
bo
. Title page to Volume 3. (put in Binder 1).
3. List of Issues published as of June 1, 1973. (put in Binder 1).
4. Replacement Pages for Tectarius (8 pages of corrections for “The Family
Littorinidae in the Indo-Pacific. Part I.” Put these 4 sheets in their proper
place in Binder | by following the looseleaf page numbers at the bottom
of the page (i.e. [05-443]). Remove and place the obsolete pages at the
end of Binder 3, just after the guide tab “Replaced Pages.”
This instruction sheet, issued June 1, 1973, may
be destroyed, or placed at the end of Binder 3.
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
Gabrielona 4)
THE GENUS GABRIELONA (PHASIANELLIDAE) IN THE
INDO-PACIFIC AND WEST INDIES
by RosBert ROBERTSON
Pilsbry Chair of Malacology
The Academy of Natural Sciences of Philadelphia
This is the first part to be published of a mono-
graph on the systematics of all the Indo-Pacific
Phasianellidae. The other parts will comprise a
general introduction, methods, a_ bibliography,
index, and treatments of the genera Phasianella
and Tricolia (including the subgenus Hiloa Pils-
bry). Conventional approaches to the systematics
of these groups, i.e. studies of shells and radulae,
have yielded inconclusive results on how species
should be categorized, and further biological
studies are planned on them. The data on Gab-
rielona systematics are published in the mean-
time because they involve fewer unresolved prob-
lems and because there are new taxa (G. pisinna,
G. raunana goubini and G. sulcifera). Biological
data relevant to systematics are unlikely to be
obtained for this rarely collected genus.
Small, low-spired Tricolia specimens, which are
far more common than Gabrielona, have been
mistaken for Gabrielona. The generic characters
detailed in this paper should help to dispel such
misidentifications in the future. Gabrielona is
distantly related to the other two genera grouped
in the Phasianellidae.
Genus Gabrielona Iredale, 1917
Type-species: Phasianella nepeanensis
Gatliff and Gabriel, 1908
Chief distinguishing characters—The shells are
small (G. hadra, the largest known species, can
be 3.3 mm. long), with globose outlines and low
Plate 36. Gabrielona nepeanensis (Gatliff and Gabriel). Figs.
1:2, 5, 7-9. South Australia. Figs. 3-4, 6, 10. Victoria, Austra-
lia. Fig. 10. Holotype. Fig. 1, x30; Figs. 2-10, x20.
[04-151]
42. Gabrielona
spires. Sculpture, if present, is most prominent
on the early whorls. Colors are pinks, reds, or
browns, and white. Inside the aperture there is a
spiral palatal sulcus which, if high in the aper-
ture, is positioned beneath a spiral, subsutural
feature of the color pattern on the external sur-
face. All the species but G. raunana have from
one to three apertural denticles. All the species
are umbilicate, with an umbilical channel.
From shells of other Phasianellidae of generally
similar shape and size, Gabrielona is most read-
ily recognized by its operculum which, external-
ly, has a pronounced marginal ridge and a con-
cave central region. The other two genera have
opercula that, externally, are convex. When fully
withdrawn into the aperture, the operculum of
Gabrielona fits shallowly against an axial ridge
(or series of faint ridges); the opercula of the
other genera fit more deeply against similar, but
generally faint, ridges. The axial ridge of Gab-
rielona very rarely is as deep in the aperture as
shown in Pl. 52, fig. 3, ar. Gabrielona radulae
are highly distinctive.
Taxonomic history—When Iredale proposed
Gabrielona, he did “not think it has really any
close relationship with Phasianella,” even though
suggesting that it “may be classed for the pres-
ent in the family Phasianellidae,” and “judging
from the shell characters and the operculum and
dead animal of the [unnamed] Lord Howe spe-
cies ... a Naticoid affinity suggests itself.”
Characteristically, Iredale did not describe the
genus or mention any distinguishing characters
other than the naticoid operculum.
Retaining Gabrielona in the subfamily Phas-
ianellinae, Thiele (1929) stated it to be doubt-
fully distinct from Chromotis H. and A. Adams,
1863. Following their description of what later
came to be the type-species of Gabrielona, Gat-
liff and Gabriel (1908) had mentioned a similar-
ity to Phasianella neritina Dunker—the type-spe-
cies of Chromotis. Wenz (1938) retained Gabriel-
ona as a tentative synonym of Chromotis, which
he ranked as a subgenus of Tricolia. Australian
malacologists have disregarded Thiele and Wenz,
and have followed Iredale in treating Gabrielona
as a distinct genus, but without giving reasons.
Finding that Gabrielona definitely is not a syn-
onym of Chromotis (here considered a synonym
of Tricolia), I re-established it, accorded it gener-
ic rank, and described it for the first time (Rob-
ertson, 1958). My study of the radula of “G.
brevis (Orbigny)” [= G. sulcifera Robertson]—
not the type species—revealed that it is rhipido-
glossate but different from that in other Phasi-
Robert Robertson
Phasianellidae
anellidae. Thus, Gabrielona does not belong in
the Naticidae or any other group with taenio-
glossate radulae. Absence of a nacreous internal
shell layer, and possession of a calcareous oper-
culum necessitated my retaining Gabrielona in
Phasianellidae, where I arbitrarily placed it in
the subfamily Tricoliinae.
Relationships— The relationships of Gabrielona
to other supraspecific taxa in the Phasianellidae
are obscure. The resemblances with most other
phasianellids—shells of small size with bright col-
ors and complex and variable patterns—seem
superficial. In view of the distinct apertural, op-
ercular and radular characters, the genus may
not even belong in the family. However, pending
more thorough knowledge of relationships within
the Trochacea, it seems best to continue to retain
Gabrielona in the Phasianellidae.
Shell description (abbreviations refer to the
drawings)—Attains lengths of 1.1-3.3 mm., widths
of 1.1-3.1 mm., and 3.0-4.6 whorls; spire angles
90°-125°; outlines globose; fairly thin to thick,
and transparent to opaque. Protoconchs (p) in-
sert to slightly exsert, slightly inflated, smooth or
with a spiral keel, white, either not demarcated
from teleoconch or slightly to fairly prominently
demarcated, 0.9-1.2 whorls when demarcated.
First whorls 0.20-0.29 mm. in diameter. Subse-
quent whorls: inflated, rounded in profile; su-
ture slightly impressed. Sculpture predominantly
smooth, or with axial plicae or spiral keels, cords
and sulci; if present, sculpture most pronounced
on second whorl, gradually declining and lacking
near periphery or wholly lacking on last whorl of
large shells. Periostracum not observed, but very
thin layer possibly present. Colors: pinks, reds,
or browns, and white. Patterns various, but com-
monly with subsutural, subperipheral, and _peri-
umbilical discordances. Aperture roundly pyri-
form; outer lip never thickened or everted, thin
to thick; callus on upper parietal area thin to
thick; a faint to fairly prominent axial ridge (ar)
or series of faint ridges within the aperture
(against which the fully withdrawn operculum
abuts); the ridges fairly variable in position, but
never distant from the edge of the outer lip; a
fairly faint to fairly pronounced spiral palatal sul-
cus (ps) high in aperture or near middle, com-
monly positioned beneath a color pattern feature
on external surface and terminating at apertural
ridge. Depending on the species, 0-3 faint to
rather prominent apertural denticles present,
namely 1 palatal and opposite the shoulder in
position (pd), 1 on the lower part of the colu-
mella (cd,)—both these on the apertural ridge—
[04-152]
May 30, 1973
and 1 on the middle part of the columellar lip
(cd,); in species with a lower columellar denticle
(cd,), a faint, broad ridge spirals up the columel-
la; apertural ridge, palatal sulcus, and apertural
denticles fairly faint to lacking on small shells.
Channel extending into umbilicus (uc) bordered
on right by the outer edge of the columellar lip
(a slope or an escarpment), and on the left (at an
acute angle) by a fairly faint to prominent es-
carpment arising from the outer edge of the
lower half of the columellar lip (abnormally, es-
carpment absent); umbilicus narrow to wide.
Opercula— Unlike those of the two other genera
in the Phasianellidae, the opercula of Gabrielona
are externally concave. In view of the extreme
rarity of live-collected Gabrielona in collections,
it is convenient that these distinctive opercula
are retained—albeit infrequently—in the apertures
of some empty shells. The fresh opercula are
fairly transparent and white. They all are pauci-
spiral and have fine, irregular spiral and radial
growth lines and wrinkles on the external and in-
ternal surfaces; this fine sculpture is not shown
in the accompanying drawings except where spe-
cially prominent.
The external surface has a prominent spiral
ridge near the edge (except the non-spiral colu-
mellar sector) and a flattish but slightly concave
central region which can have a differently tex-
tured or sculptured outer, spiral area. On all
Gabrielona opercula, a callus of varied thickness
overlays the upper two-thirds or three-quarters of
the central region. This callus covers all or most
of the spiral suture, is thickest near the middle
columellar edge of the operculum, and its lower
margin (marked c¢ on the drawings) is irregular
and variable in position and height, with or with-
out a distinct escarpment. Small opercula and the
single one known of G. sulcifera have a thin or
very thin callus and therefore the suture is clear-
ly visible externally—as well as in transparency.
In the different species, the spiral ridge and cen-
tral region are variously sculptured. The non-
spiral columellar edge of the operculum is bev-
eled.
The internal surface of the operculum is rela-
tively flat, but the central part of the last whorl
near the columellar margin can be slightly con-
cave. The early whorls are slightly raised, and a
low, spiral escarpment is at the suture. The cen-
tral area (at the axis) on all the species with op-
ercula available (G. nepeanensis, G. pisinna, G.
raunana [both subspecies] and G. sulcifera) is
circular, crested at the perimeter, and 0.11-0.14
mm. in diameter—i.e., correlates neither with the
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
Gabrielona 43
various diameters of the first whorl of the shell,
nor with the maximum shell size attained by the
various species. This circular central area can be
conspicuously smoother, shinier, and more trans-
parent than the remainder of the internal sur-
face, and always bears a low, central boss. The
operculum of G. raunana (both subspecies) dif-
fers from those of the other species in having
structural radial lamellae near the edge. Muscle
attachment scars are never clearly defined.
When fully withdrawn into the aperture, the
operculum abuts against the apertural ridge (ar),
and fits closely against the columellar lip as
well as the inner surface of the outer lip. In
those species possessing the two columellar den-
ticles, the operculum pivots against the columel-
lar lip between these. The slight differences be-
tween species in the form of the columellar lip
account for the slight differences in the outline
of the columellar margin of the opercula.
Anatomy—The very few live-collected speci-
mens available all were preserved dry, so that
the bodies were dried out and appressed against
the outer shell wall in the aperture. In G. pis-
inna, the gut—distended with whitish, calcareous
fragments—was conspicuous, fairly short and U-
shaped, arising at the left, extending posteriad
and bending to the right at a position about half
of a whorl back from the outer lip, and then ex-
tending forward to the anus on the right near the
outer lip. There was no indication of an enlarged
stomach. The mantle edge appeared to be rather
thick, but the total volume of the dried body
seemed remarkably small. Jaws were not detect-
ed but could be present.
Radulae (Pls. 42 and 57)—I have been able to
study the radulae of only G. pisinna and G. sul-
cifera. These differ greatly from other known
phasianellid radulae and also from each other.
The specimens from which the radulae of the
two species were extracted were collected about
9,200 miles apart (New Caledonia and Antigua),
and the size difference between their shells is
considerable: the shell volume of G. pisinna is
about 19-fold smaller than that of G. sulcifera.
Relative to shell size, the radula of G. pisinna is
rather longer and considerably wider than that of
G. sulcifera. These differences in actual and rela-
tive sizes perhaps account for some of the great
differences. In particular, the small absolute size
(but not the greater relative width) of the radula
in G. pisinna could account for the fewer margin-
als and laterals (3 laterals instead of the 5 in G.
sulcifera), and the more curved transverse rows
of teeth. The “central” of G. pisinna possibly is a
[04-153]
44 Gabrielona
pseudocentral comprised of one or two pairs of
the original innermost laterals fused together
(perhaps also with the original central).
The size differences could not account directly
for some of the other interspecific differences:
the multicusped laterals of G. pisinna and, in G.
sulcifera, the unwinged central and laterals, the
differently-shaped bases of the laterals, and the
massive innermost marginals.
The radular differences help to confirm the
conclusion reached independently from study of
the shells and opercula of the two species, name-
ly that they are distantly related congeners. Lack-
ing information as to the total diversity of Gab-
rielona radulae, their possible ontogenetic
changes, the structural consequences of their ab-
solute and relative sizes, and bearing in mind the
evolutionary plasticity of Tricolia radulae, I con-
sider the radular differences inadequate evidence
for separating G. sulcifera from G. pisinna in a
different subgenus or genus.
Relationships within genus—Related pairs of
Recent taxa are all allopatric. Only in New Cal-
edonia is more than one species of Gabrielona
known to occur, and these (G. pisinna and G.
raunana) are distantly related congeners.
The most closely related taxa distinguished
here are G. raunana raunana and G. raunana
goubini, which are ranked as subspecies. G. pis-
inna is a dwarf, tropical Indo-Pacific homologue
of southeastern Australian G. nepeanensis. G.
sulcifera of the Caribbean perhaps is related, al-
beit fairly distantly, to the western Pacific G.
raunana. G. hadra is a clearcut fossil precursor
of G. sulcifera. G. nepeanensis and G. pisinna
seem distantly related to their congeners.
Color and pattern variations—All the species
are variable in coloration and pattern: G. rau-
nana and G. nepeanensis especially so, and G.
pisinna least of all. The range of coloration of
Gabrieclona is narrower than in each of the other
two phasianellid genera. Most of the color and
pattern variation in Gabrielona is gradational,
but a discontinuous color variation is treated
under G. raunana goubini.
Sexual dimorphism—Not detected conchological-
ly.
Fossil history—The only fossil species known
certainly to belong in the genus is “Tricolia”
hadra Woodring from the Bowden Formation
(Middle Miocene or possibly Pliocene-Pleisto-
cene) in Jamaica, West Indies.
Distribution of Recent species—Until I transfer-
red a West Indian species to the genus (Robert-
son, 1958), G. nepeanensis (Gatliff and Gabriel),
Robert Robertson
Phasianellidae
from southeastern Australia, was the only named
species in the genus. An unnamed and cursorily
studied Gabrielona, mentioned by Iredale (1917),
was live-collected in the “sub-littoral” at Lord
Howe Island (about 400 miles east of the coast
of northern New South Wales, Australia). This
locality, shown with a circle on Pl. 39, is be-
tween the known distributions of G. nepeanensis
and the two tropical Indo-Pacific species. Ire-
dale’s specimens from Lord Howe were not lo-
cated at the Australian Museum by Dr. D. F.
McMichael in 1962 (letter to Dr. R. T. Abbott
dated July 10).
One new tropical Indo-Pacific species, G. pis-
inna, and one new subspecies, G. raunana gou-
bini, are described and named here, and the
West Indian species to which I misapplied the
name Phasianella brevis Orbigny is named G.
sulcifera.
Gabrielona quite possibly occurs in other trop-
ical and subtropical faunal areas, such as West
Africa and the Panamic Province. The four
known Recent species occur almost exclusively
along the coasts of continents and high islands;
the only known exception is G. raunana raunana
Ladd, a subspecies known only as subfossil shells
from an atoll.
Abundance—Very few live-collected specimens
of any of the species have been available for
study: only 7 G. pisinna and 1 G. sulcifera. The
other two Recent species, including the type-
species G. nepeanensis, are known only from
empty, beach worn or subfossil shells, a few with
opercula. Thus, the genus appears to be a relict
group. Alternatively, it may have been rarely col-
lected if, as I suspect, it mainly lives well below
the tidal zone in algae on rocks.
Habitats—Known only from among algae in
shallow water (G. pisinna) or in sand (G. sulci-
fera); for details see under these species. G. pis-
inna may live as deeply as 8 fathoms; a probably
adventitious shell of G. sulcifera came from 287
fathoms.
Larval ecology—The fairly small range of varia-
tion in the diameter of the first whorls of the six
known taxa of Gabrielona (0.20-0.29 mm.) and
the small sizes presumably indicate that the full-
grown larval shells are small and relatively uni-
form in size, and that the larvae are all pelagic
and planktotrophic.
Abnormalities—A_ striking series of abnormal-
ities possibly caused by an individual living in an
unusual habitat is described under G. sulcifera.
Abnormal growth caused by incrustations are
rare in Gabrielona; one case is reported under G.
nepeanensis.
[04-154]
May 30, 1973
Synonymy-
1917 Gabrielona Iredale, Proc. Malac. Soc. London 12:322
[listed], 327. Type-species (by monotypy): Phasianella
nepeanensis Gatliff and Gabriel, 1908.—1929, Thiele,
Handb. syst. Weichtierkunde, Jena, 1:70.—1938, Wenz,
Handb. Palaozool., Berlin, 6(1)Teil 2 [Prosobranchia],
p. 362,— 1958, Robertson, Johnsonia 3(37):246-260.
Excluded species—In 1958 (pp. 253, 257), I sug-
gested that three American Miocene (or Plio-
Pleistocene)species might belong in Gabrielona:
Tricolia (Eulithidium) hadra Woodring, Didia-
nema P waltonia Gardner, and Tricolia P syn-
toma Woodring. T. hadra is here referred defi-
nitely to Gabrielona, but subsequent study of the
holotypes of the other two species has shown
that neither belongs in the Phasianellidae. Two
upper Tertiary species from northern Venezuela
described as Gabrielona are treated under Ex-
cluded Species on p. 61.
The possible second Western Atlantic Recent
species of Gabrielona (Robertson, 1958, p. 259),
from Brasil, proves upon restudy to be a depau-
perate, low-spired Tricolia.
Gabrielona nepeanensis
(Gatliff and Gabriel, 1908)
(Pls. 36-39)
Range— Recent: known only from South Austra-
lia and Victoria, Australia. Possibly occurs also
at Tasmania, but not yet known there.
Chief distinguishing characters—The shell dif-
fers from those of all other known species in the
genus except G. pisinna in having both columel-
lar denticles (ed; and ed,), in lacking axial or
spiral sculpture, and in opercular characters (see
under G. raunana and G. sulcifera for differ-
ences). Differs from all other species, including
G. pisinna, by its complex and varying but con-
sistently distinct color patterns (the adults always
with a colorless spiral band opposite the palatal
sulcus). For detailed differences from G. pisinna,
the most similar species, see under that species.
Abundance—36 shells from beach sand avail-
able, 1 with operculum in place in aperture; none
live-collected (probably lives below the tidal
zone).
Abnormal shell—One shell (PI. 36, fig. 6) has a
double outer lip near the suture, a columellar
callus slightly detached from the palatal area,
and the upper columellar denticle (cd,) fainter
than usual. Some incrustation must have inter-
fered with normal growth.
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
Gabriclona 45
Shell description—Attains length of 1.9 mm.,
width of 1.7 mm., and 3.9 whorls; spire angle
95°-105°; length invariably equals or exceeds
width (except for small shells); fairly thick but
slightly translucent to fairly transparent. Proto-
conch insert, slightly inflated, smooth, white, not
demarcated from teleoconch. First whorl 0.24-
0.28 mm. in diameter. Penultimate and _ last
whorls: slight flattening below suture but no dis-
tinct shoulder (PI. 38, fig. 1); surface smooth ex-
cept for axial growth lines and slight wrinkles,
fairly shiny. Colors: pale to dark pink (rarely,
tinged with orange), and white. Patterns: alter-
nating pink and white subsutural marks, each
becoming divided by a virtually colorless spiral
band directly opposite the palatal sulcus; 9-12
paired pink marks on last whorl; predominant
below subsutural area: axially aligned wavy pink
stripes or irregular marks; subperipheral series
fairly faint pink and white marks; inner umbilical
area colorless, surrounded by spiral series short
white axial stripes (commonly, partially coa-
lesced); rarely, zigzag pale pink ‘stripes or irregu-
lar marks entirely replace usual pattern (PI. 36,
figs. 3-4). Outer lip and callus on upper parietal
area fairly thin; no palatal denticle; palatal sul-
cus high in aperture; both columellar denticles
present, most prominent on large shells (PI. 38,
fig. 2). Columellar lip thickened adjacent to cen-
tral part umbilical channel, and an escarpment
present; escarpment to left of umbilical channel
fairly faint to prominent, arising fairly low off
the outer edge of columellar lip; umbilicus nar-
row to fairly wide.
Shell measurements (mm.)—
length width no. whorls
1.92 1.66 3.9 large; South Australia
1.56 1.46 3.3 average; South Australia
0.80 0.87 2D smallest; South Australia
Plate 37. Gabrielona nepeanensis (Gathff and Gabriel). Vic-
toria, Australia. Original figures of holotype (from Gatliff
and Gabriel, 1908, pl. 21, figs. 9-10), enlarged. Both x20.
Fig. | incorrectly shows a prominent shoulder (compare PI.
36, fig. 10).
[04-155]
46 Gabrielona
Plate 38. Gabrielona nepeanensis (Gatliff and Gabriel). Fig.
1. Outline of large shell, x17. Fig. 2. Aperture, x33; ar, aper-
tural ridge; ed, and edz, upper (outer) and lower (inner) col-
umellar denticles (respectively); ps, palatal sulcus; uc, umbil-
ical channel. Figs. 3-5. Operculum. Internal surface, longitu-
dinal section, and external surface (respectively). All x33.
Operculum-(Pl. 38, figs. 3-5). Only a single,
abraded operculum of this species has been
available, and it has been difficult to determine
the sculpture of its external surface. The crest of
the spiral ridge near the edge seems to be round-
ed, and on the central region—between the outer,
spiral area and the central area—there is a low,
spiral ridge extending to a prominence on the
non-spiral columellar margin. The outline of the
columellar margin at the junction of the spiral
and non-spiral sectors is shallowly concave.
Otherwise the operculum of G. nepeanensis
seems to be like that of G. pisinna, only larger;
both are fairly thin.
Synonymy—
1908 Phasianella nepeanensis Gatliff and Gabriel, Proc. Roy.
Soc. Victoria, n.s., 21(1): 366, pl. 21, figs. 9-10 [re-
produced here, PI. 37] (Flinders, Western Port; Ocean
Beach, near Point Nepean [both Victoria, Australia]).
—1917, Iredale, Proc. Malac. Soc. London 12(6): 322
[listed], 327 [the only named species included in Gab-
rielona but not formally transferred].
1938 Gabrielona nepeanensis (Gatliff and Gabriel). Cotton
and Godfrey, Malac. Soc. South Australia Publ. 1: 9;
1945, Cotton, Trans. Roy Soc. South Australia 69(1):
165; 1958, Robertson, Johnsonia 3(37): 257, pl. 137,
figs. 2-3, pl. 140, fig. 1; 1959, Cotton, South Australian
Mollusca, Archaeogastropoda, Adelaide, pp. 270-271,
fig. 185 [shell shape highly inaccurate], p. 347 [listed].
Robert Robertson
Phasianellidae
Types—The holotype of Phasianella nepeanensis
Gatliff and Gabriel (Pl. 36, fig. 10; Pl. 37) is now
at the National Museum of Victoria (no. F543),
Melbourne, Australia. A small paratype is at the
Australian Museum (no. C.45057), Sydney (Rob-
ertson, 1958, pl. 140, fig. 1). Gatliff and Gabriel
did not mention the number of specimens avail-
able to them, and recorded the species from two
localities 23. miles apart. Cotton and Godfrey
(1938) selected “Flinders, Victoria” as the type-
locality. However, in 1945 and_ subsequently,
Cotton has stated that the type-locality is “near
Point Nepean,” whence (judging by the specific
name) the holotype came.
Locality records (see map, Pl. 39; literature and uncertain
records circled—SOUTH AUSTRALIA: Port Lincoln (“appar-
ently nepeanensis,” Iredale, 1917; Cotton, 1945); Mouth of
Middle River, N. coast Kangaroo I. (from beach sand, 1954,
B. Daily, via M. F. Glaessner and G. L. Harrington); Port
Adelaide (Calvert Coll., both ANSP); Robe (Cotton, 1945).
VICTORIA: Port Fairy (in beach sand, 1923, H. A. Pilsbry,
ANSP); ocean beach near Point Nepean [38 mi. S.S.W. of
Melbourne] (Gatliff and Gabriel, 1908; Natl. Mus. Vict.:
Austral. Mus.); Flinders [47 mi S. of Melbourne], Western
Port (Gatliff and Gabriel, 1908).
All three of the circled locality records in South
Australia are questionable because Cotton (1945,
1959) mentioned having difficulty distinguishing
G. nepeanensis from “Pellax virgo,” and the
locality data with all the available specimens ex-
cept one juvenile shell from Kangaroo Island
seem not wholly reliable either. A large series
(30 shells) is from the Calvert Collection (col-
lector not recorded), labeled “Pt. Adelaide” is
suspect because this is the largest port in the
area.
Gabrielona pisinna Robertson, new species
(Pls. 39-42)
Range—Recent: known only from Mauritius, In-
dian Ocean, and New Caledonia, eastern Melan-
esia. Perhaps widespread in the tropical Indo-
Pacific around high islands.
Chief distinguishing characters—The shell, usual-
ly less than 1 mm. in length, is full-grown at a
smaller size than in any other known phasianel-
lid. It resembles young G. nepeanensis in size
and number of whorls. (Further resemblances: has
both columellar denticles, an unsculptured sur-
face, and a similar operculum.) That G. pisinna
is full-grown at a smaller size than G. nepean-
ensis is shown by the complete development on
the larger shells of the lower columellar denticle
(ed,). Further differences from G. nepeanensis:
first whorl smaller; spire lower (except for some
[04-156]
May 30, 1973
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Plate 39. Geographical distributions and records of Gabriel-
ona nepeanensis (Gatliff and Gabriel), Gabrielona species?
(Iredale, 1917), and Gabrielona pisinna Robertson, which
large shells, the width of G. pisinna invariably
exceeds the length); aperture smaller than in
young G. nepeanensis of comparable size (com-
pare Pl. 40 with Pl. 36, fig. 1); color patterns dif-
ferent (finer), and reddish coloration darker.
Relationships—A dwarf, tropical homologue of
G. nepeanensis, which might need to be ranked
as a subspecies if the form occurring at Lord
Howe Island (map, Pl. 39) is intermediate (see
p. 44).
Abundance—68 specimens available: 2 empty
shells from Mauritius (1 with operculum), and 66
specimens from New Caledonia (10 with oper-
cula, 7 of these live-collected).
Habitat—All 7 of the live-collected specimens
came from washings from algae collected in 0-3
ft. on Récif Ricaudy, New Caledonia, on rocks
near shore on the fringing reef. An empty but
fresh shell with 4 drill holes and an operculum in
place in the aperture came from a depth of 8
fathoms at Mauritius (see under Locality rec-
ords). (Many of the shells have these drill holes.)
Shell description—Attains length of 1.1 mm.,
width of 1.1 mm., and 3.0 whorls; spire angle
115°-125°; width exceeds length (excepting
some large shells); fairly thin and translucent to
transparent. Protoconch insert, slightly inflated,
smooth, white, not demarcated from teleoconch.
First whorl 0.20-0.24 mm. in diameter. Penulti-
mate and last whorls: slight flattening below su-
ture but no distinct shoulder (Pl. 41, figs. 2-3);
surface smooth except for axial growth lines and
slight wrinkles, shiny; fine spiral sulci on base of
small shells (Pl. 41; fig. 1). Colors: dark pink to
bright red (rarely, tinged with amber [faded?]),
and white. Patterns: white spiral band below the
suture, coalescing with variably-shaped white
subsutural patches (7-9 on last whorl), lower
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
Gabrielona 47
*) MARIANA Nokes
* Guam
: Sopon «
oO cof
NEW
ZEALAND
1
160° 160°
G, NEPEANENSIS
perhaps is widespread in the tropical Indo-Pacific but which
presently is known only from the two areas 7300 miles apart.
edges of which are slightly below or directly op-
posite the palatal sulcus; pink or red marks or
non-axial stripes alternate with white subsutural
patches; predominant below subsutural area:
fairly regular axial or very steeply ascending
dark stripes; subperipheral series of white marks
alternating with non-axial dark stripes or axially
paired marks; umbilical area closely surrounded
by spiral series of short white axial stripes (fair-
ly commonly, partially coalesced); very rarely,
subsutural patches and subperipheral white
marks coalesce into irregular axial white bands
(Pl. 40, fig. 2). The aperture averages relatively
Plate 40. Gabrielona pisinna Robertson. Fig. 1. Mauritius.
Figs. 2-6. New Caledonia. Figs. 4-6. Holotype. All x30.
[04-157]
48 Gabrielona
Robert Robertson
Phasianellidae
Plate 41. Gabrielona pisinna Robertson. Fig. 1. Smallest spec-
imen, x60. Figs 2-3. Outlines of largest shells, both x33. Fig.
2. Unusually high-spired shell. Fig. 4. Aperture, x60; ar, aper-
tural ridge; ed, and cd, upper (outer) and lower (inner) col-
umellar denticles (respectively); ps, palatal sulcus; uc, um-
bilical channel. Figs. 5-7. Operculum. Internal surface, longi-
tudinal section, and external surface (respectively); ¢, lower
edge of callus. All x60.
slightly wider than in G. nepeanensis. Outer lip
and callus on upper parietal area thin; no palatal
denticle; palatal sulcus high in aperture, slightly
variable in position (commonly, fairly faint); both
columellar denticles present, most prominent on
large shells (Pl. 41, fig. 4). Columellar lip slightly
thickened adjacent to central part umbilical
channel, and an escarpment, when present, faint;
escarpment to left of umbilical channel fairly
prominent, arising fairly low off outer edge col-
umellar lip; umbilicus fairly wide.
Plate 42. Gabrielona pisinna Robertson. Half of one trans-
verse row of radular teeth, showing (from left to right) the
5-cusped central (or pseudocentral?), the three laterals on the
right side, and the entire row of marginals. The cusps of the
central and laterals are stippled. New Caledonia. x1050.
Shell measurements (mm. )—
length width no. whorls
1.09 1.10 3.0 largest; New Caledonia
0.91 0.93 2.8 holotype; New Caledonia
0.70 0.75 2.6 average; Mauritius
0.34 0.42 1.7 smallest (Pl. 41, fig. 1);
New Caledonia
Operculum—(PI. 41, figs. 5-7)- The spiral ridge
near the edge on the external surface of the fair-
ly thin operculum is sharply crested except near
the columellar margin where it is rounded. The
inner margin of the ridge is angled or rounded.
The central region has an outer, spiral area with
a mat surface contrasting with a shiny, more
transparent central area. The single operculum
available from Mauritius differs from those from
New Caledonia in having a faint spiral ridge be-
tween the two areas (like that of G. nepeanen-
sis). Fairly prominent wrinkles parallel to the
non-spiral columellar margin are present to the
left of the central part of the callus. The lower
margin of the callus (c) varies in position.
Radula (Pl. 42; 2 studied)—Attains length of
0.37 mm., width of 0.08 mm., and with as many
as 20 very strongly curved transverse rows of
teeth (including a few nascent rows). Central (or
pseudocentral?) 5-cusped, laterally winged ante-
riorly, and with posterolateral projections on the
base. Three laterals, each with long pointed
cusps (the two innermost generally with 5 cusps,
and the outermost with 6); the two outermost
laterals have the largest distal portions, and the
outer lateral is positioned posterior (rather than
lateral) to the morphologically middle lateral; the
[04-158]
May 30, 1973
base of the innermost lateral bears a posterolat-
eral peg that fits into the middle of the inner
edge of the base of the middle lateral, and the
two outermost laterals have large bases that ex-
tend posterolaterally beneath the marginals. As
many as 13 pairs of marginals (lowest count 10),
each row widely overlapping the row posteriad.
All the marginals have elongate distal portions,
and the innermost of these are serrate on the
outer (posterior) edge; the serrations become fin-
er outwards and are absent altogether on the
small outermost teeth.
Types—The holotype (Pl. 40, figs. 4-6), from
Récif Ricaudy, near Noumea, New Caledonia, is
at the Academy of Natural Sciences of Philadel-
phia (no. 301611). So also are paratypes from
the type-locality (no. 271062) and from other lo-
calities: Récif de Gatope, New Caledonia (nos.
267567 and 267568), and Mauritius (nos. 273188
and 273328). Paratypes from New Caledonia will
be distributed to USNM, MCZ, BM, and IrSnB
(2 shells each).
Derivation of new name—Latin, pisinnus, little.
Locality records (see map, Pl. 39)—MAURITIUS: % mi.
N.N.E. of Flic en Flacq Pt. (1-10 ft., from Caulerpa wash-
ings, Nov., R.E.M. Ostheimer & V. Orr Maes, Sta. M 203):
Black River Bay, 1 mi. W.N.W. of mouth Black River, both
W. coast (dredged 8 fms. [1 empty but fresh shell], coarse
sand, broken shell, very little weed, Nov. 5, both 1960,
R.E.M. Ostheimer, J. de B. Baissac & V. Orr Maes, Sta. M
208, both ANSP). NEW CALEDONIA: Grand Reéif de Ga-
tope, 7% mi. W. of Voh (dredged 6-18 ft., inner edge of bar-
rier reef, sand, weed, coral rubble, Dec. 31, 1960 & Jan. 2,
1961, Stas. K 538 & K 539); E. end Reécif Ricaudy, 2% mi.
S.S.E. of Noumea (0-3 ft., Jan. 11, 1961, both G. & M. Kline
& V. Orr Maes, Sta. K 553, both ANSP).
Gabrielona raunana Ladd, 1966
(Pls. 43-50)
Range—Recent: known only from Eniwetok (an
atoll), northwestern Marshall Islands (G. raunana
raunana Ladd), and from the Loyalty Islands and
New Caledonia (all high islands), eastern Melan-
esia (G. raunana goubini Robertson).
Chief distinguishing characters—This is the only
species in the Phasianellidae having a shell with
prominent axial sculpture. The plicae are most
prominent and regularly arranged on the second
whorl, and are obscure or absent at and near the
periphery of the last whorl of large shells. Also
differs from other species in the genus as fol-
lows: spiral keel on protoconch; all apertural
denticles lacking; operculum with obliquely radi-
al sulci on part of external surface, and with
INDO-PACIFIC MOLLUSCA vol. 3, no. 14
Gabrielona 49
structural radial lamellae at and near the spiral
outer edge. The outlines of the shells and color
patterns of both subspecies are distinct from
each other and from those of the other species.
Remarks—No live-collected specimens are avail-
able of either subspecies; most of the shells are
beach worn or subfossil. G. raunana, known only
from populations in two areas slightly more than
2,000 miles apart, seems thus to be a relict spe-
cies.
Shell descriptions, Types, Locality records, etc.
See under G. raunana raunana and G. raunana
goubini.
Operculum (Pl. 44)—Only 2 opercula of G.
raunana raunana and 8 of G. raunana goubini
are available, and because all these are abraded
or corroded—making description and illustration
of the original sculpture difficult—and because it
is doubtful whether the seeming slight differ-
ences between the two subspecies are real, they
are discussed together here. The operculum in
best condition is a small one from G. raunana
raunana.
Relative to those of G. nepeanensis and G.
pisinna, the operculum of G. raunana (both sub-
species) is thick and comprises fractionally more
whorls (comparing opercula of the same size).
The external surface has a central region with an
outer, spiral area distinctive in having prominent,
obliquely radial sulci (irregular and varying in
spacing), and a central, smoothish area with a
fairly thick callus. The crest of the spiral ridge is
Plate 43. Gabrielona raunana Ladd. Thinly coated with mag-
nesium oxide to accentuate the sculpture and obscure the
color pattern. Figs. 1-2. G. raunana raunana Ladd. Eniwetok,
Marshall Islands. Figs. 3-4. G. raunana goubini Robertson.
Lifou, Loyalty Islands. All x20.
[04-159]
50 Gabriclona
Phasianellidae
Robert Robertson
near the outer edge, and (at least on G. raun-
ana raunana) its whole surface is irregularly
wrinkled and knobbed, the wrinkles tending to
be aligned with the adjacent obliquely radial
sulci.
The erosion of the external surface makes sev-
eral features obscure. The spiral ridge is strongly
abraded on all the opercula available from G.
raunana goubini. On all but one of these there is
a deep but irregular central pit that may be an
erosional feature; this is margined by an irregu-
lar but steep escarpment at the lower margin of
the callus (c). On the unpitted operculum of G.
raunana goubini, two spiral sulci are near the
center and there is no steep escarpment border-
ing the callus; this perhaps is the uneroded orig-
inal sculpture.
Seen in transparency, near the outer edge of
the operculum are structural radial lamellae at
right angles to the horizontal plane of the oper-
culum. These are closely and regularly spaced
and extend a uniform distance from the edge.
Most prominent at the edge of the operculum of
G. raunana raunana, these project from the sur-
face as external lamellae, thus causing the out-
line to be finely notched. These external lamellae
perhaps are erosional features.
Gabrielona raunana raunana Ladd, 1966
(Pl. 43, figs. 1-2; Pl. 44, figs. 4-6; Pls. 45-47)
Range—Recent: known only from subsurface de-
posits on Eniwetok Atoll, northwestern Marshall
Islands (obtained from drillings). Perhaps wide-
spread at atolls in Micronesia or the whole tropi-
cal northwest Pacific.
Chief distinguishing characters—The shell of
this subspecies has fairly regularly spaced but
commonly coalesced colorless or white spots,
each one surrounded by 6 others. More differ-
ences are given under G. raunana goubini. Sculp-
ture is almost identical in the two subspecies.
Abundance—32 subfossil shells available, 1 with
an operculum in place in the aperture; 1 loose
operculum.
Shell description—Attains length of 2.1 mm.,
width of 2.1 mm., and 3.4 whorls; spire angle
110°-125°; width invariably equals or exceeds
length; fairly thick to fairly thin and opaque to
fairly transparent. Protoconch insert, smooth ex-
cept for slightly descending spiral keel, white,
slightly demarcated from teleoconch, 0.9 whorl.
First whorl 0.26-0.29 mm. in diameter. Penulti-
mate and last whorls: flattening below suture,
and faint shoulder (Pl. 46, fig. 1). Sculpture of
second whorl: 18-30 strong axial plicae, convex
towards outer lip, highest, most sharply crested
and most widely spaced on first half where a fairly
prominent to fairly obscure spiral cord extends
from the keel on protoconch. On later whorls pli-
cae smaller, crests rounded, more closely spaced,
and less regular in arrangement and _ structure,
commonly with intercalated secondary plicae be-
low suture and with divarications and anasto-
moses below periphery. A faint spiral cord sur-
rounds the umbilical area of young shells; rarely,
faint reticulations on base of medium-sized shells.
On last whorl irregular axial plicae most promin-
ent below suture and on base, absent near peri-
phery of large shells where surface is smooth
except for axial growth lines. Surface shiny,
Colors: pale (faded?) to fairly pale pink or yel-
lowish brown, and white. Patterns: more or less
quadrate pink or brownish subsutural patches
(6-10 on last whorl), lower edges directly opposite
palatal sulcus; surface almost entirely covered
with fairly regularly spaced spots, each one sur-
Plate 44. Gabrielona raunana Ladd. Opercula. Figs. 1-3. G.
raunana goubini Robertson, x33. Figs. 4-6. G. raunana raun-
ana, x60. Internal surface, longitudinal section, and external
surface of each; c, lower edge of callus.
[04-160]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
Gabrielona 5]
Plate 45. Gabrielona raunana raunana Ladd. Subsurface Re-
cent deposits, Eniwetok Atoll, Marshall Islands. Fig. 7.
Holotype. Figs. 1-2, x40; Figs. 3-7, x20.
rounded by 6 others; spots white in pale areas
alternating with subsutural patches and colorless
elsewhere except (somewhat commonly) for sub-
peripheral spiral series and (rarely) spiral series
on base; commonly: spots irregularly coalesced;
ground color pale; fairly commonly: subperipheral
spiral series irregular, slightly darkened markings
alternating with white-spotted areas; closely sur-
rounding umbilicus: spots variously coalesced in-
to irregular, steeply descending white stripes on
colorless ground; fairly rarely: wavy, axial dark-
ened bands extending from subsutural patches to
base; rarely: almost a uniform pink with no spots
near periphery. Outer lip and callus on upper
parietal area somewhat thin to fairly thick; no
palatal denticle; palatal sulcus high in aperture,
Plate 46. Gabrielona raunana raunana Ladd. Fig. 1. Outline
of large shell, x17. Fig. 2. Aperture, x33; ar, apertural ridge,
ps, palatal sulcus; uc, umbilical channel.
neither columellar denticle present (Pl. 46, fig.
2). Columellar lip not thickened, and steep slope
into umbilical channel but no escarpment; es-
carpment to left of umbilical channel prominent,
arising quite high off the outer edge of the col-
umellar lip; umbilicus fairly narrow to wide (PI.
46, fig. 2).
Shell measurements (mm.)—
length width no. whorls
2.09 2.09 3.4 largest
1.60 1.67 3.1 average; holotype
0.89 1.07 2.4 smallest
O perculum—See under G. raunana (species).
Synonymy—
1966 Gabrielona raunana Ladd, [U.S.] Geol. Surv. Prof.
Paper 531, pp. 13 & 17 [listed], 54 [described], pl. 10,
figs. 1-5 (Recent, Eniwetok Atoll).
Types—The holotype (Pl. 45, fig. 7), out of a
drilling from 20-45 ft. below land surface Eluge-
lab, Eniwetok, is at the United States National
Museum (no. 648319), Washington, D.C. So also
are all the paratypes except 3 donated to the
Academy of Natural Sciences of Philadelphia
(no. 302131).
Locality records (see map, Pl 47)— MARSHALL ISLANDS:
Elugelab (20-60 ft. deep), Parry (30-45 ft., 90-110 ft., 1865-
1895 ft.), and Mujinkarikku (35-40% ft.), all Eniwetok Atoll
(from 8 drill holes in Recent subsurface deposits; 30 shells
and 1 loose operculum at depths of 20-60 ft., 1 shell from
90-110 ft., and 1 probably adventitious shell in lower Mio-
sene strata at 1865-1895 ft., all about 1952, IL.S. Ladd,
USNM & ANSP). For an account of the drilling operations
which yielded most of the specimens, see Ladd and Schlanger
(1960, [U.S.] Geol. Surv. Prof. Paper 260-Y, pp. i-iv, $63-
905).
[04-161]
52. Gabrielona
| g 4° ' Iwo Jima . Mores
poo SW rormosA Cs
“MARIANA
2 Is.
G. RAUNANA RAUNANA ye ye
o go"
Enwetdh@ \ 7 © MARSHALL IS.
|, Oth Sasle ‘
“Y = + +, Wot/e
AO oe Sa
Cee ee Ponape +, Majuro
ROLINE Me Se C
Ss,
4 . | See oo us
eae a
5 | New NF, ee ef
) CALEDONIA WP
a 1 G. RAUNANA GOUBINI
20° \40°
Plate 47. Geographical records of Gabrielona raunana_ rau-
nana Ladd and Gabrielona raunana goubini Robertson.
Robert Robertson
Phasianellidae
Gabrielona raunana goubini
Robertson, new subspecies
(Pl. 43, figs. 3-4; Pl. 44, figs. 1-3; Pls. 47-50)
Range—Recent: known only from Lifou, Loyalty
Islands, and Ile des Pins, New Caledonia, eastern
Melanesia. Perhaps widespread around high is-
lands in Melanesia or the whole tropical south-
west Pacific.
Chief distinguishing characters—The shell dif-
fers from that of the nominate subspecies as fol-
lows: spire averages higher and aperture rela-
tively smaller; first whorl averages smaller; pli-
cae finer on medium-sized shells; color pattern
almost invariably with steeply descending pink or
yellowish brown stripes, and white subsutural
patches commonly ring-shaped.
Abundance—1,116 shells available, 8 with an
operculum in place in the aperture. All the speci-
mens were sorted from beach sand; some are
freshly dead, but many are worn or broken and
some are bleached.
Plate 48. Gabrielona raunana goubini Robertson. Lifou, Loyalty Islands. Fig. 7. Holotype. Figs. 1-2, x40; Figs. 3-10, x20.
[04-162]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
Gabriclona — 53
Remarks—In view of the apparent abundance of
this subspecies in beach sand it is remarkable
that it has not been named previously, in spite of
rather extensive study of small marine shells
from the Loyalty Islands (Tomlin, 1936, Proc.
Malac. Soc. London 22(3): 145-152).
Color variation—Most of the shells have pink
markings; an infrequent color form is entirely de-
void of pink and has pale yellowish brown mark-
ings; intermediates have the pink followed by the
yellowish markings. Only a few shells were not
readily sorted into one of these three categories.
Of the 1,088 shells from Lifou, approximately
938 (86%) are the pink form, 112 (10%) are inter-
mediates, and 38 (4%) are the yellowish form.
The frequencies of the three color forms are
comparable in the much smaller sample (28
shells) from Ile des Pins: 18 (64%) pink, 7 (25%)
intermediates, and 3 (11%) yellowish.
Shell description—Attains length of 2.4 mm.,
width of 2.2 mm., and 3.9 whorls; spire angle
90°-115°; width fairly commonly exceeds length;
fairly thick but slightly translucent to fairly trans-
parent. Protoconch like that of G. raunana raun-
ana. First whorl 0.23-0.28 mm. in diameter. Sub-
sequent whorls: flattening below suture, and
faint to obscure shoulder (PI. 50, fig. 1). Sculp-
ture like that of G. rauwnana raunana except pli-
cae (PI. 50, fig. 2) finer, spiral cord on first quad-
rant(only) of second whorl faint or absent, and
no reticulations on base of medium-sized shells.
Surface shiny. Colors: fairly pale to dark pink,
pale yellowish brown, and white. Patterns: vari-
ably-shaped white subsutural patches (7-9 on last
whorl), commonly ring-shaped with axial stripe
extending to suture, lower edges near or direct-
ly opposite palatal sulcus; white subsutural
patches alternating with variable yellowish
brown or pinkish marks; predominant below sub-
sutural area: steeply descending pink and/or yel-
lowish brown stripes; commonly: subperipheral
series small, irregular white marks alternating
with slightly darkened pinkish or yellowish
brown marks; fairly commonly: similar series
small white marks on base, with darkened pink-
ish marks; closely surrounding umbilicus: steeply
descending, partially coalesced white — stripes:
very rarely: all descending stripes zigzag or bro-
ken into irregular marks (PI. 49). Aperture aver-
ages slightly narrower and _ relatively smaller
than that of G. raunana raunana; outer lip some-
what thick; callus on upper parietal area thin to
fairly thick; apertural denticles, palatal sulcus,
columella and umbilical area as in G. raunana
raunana, except escarpment to left of umbilical
channel fairly faint to prominent and umbilicus
narrow to quite wide.
Shell measurements (mm.)—
length width no. whorls
2.42 2.16 3.9 largest
1.92 1.80 3.5 average; holotype
0.90 1.01 2.6 smallest
Plate 49. Gabrielona raunana goubini Robertson, Rare color
patterns, Figs. 1-3. Lifou, Loyalty Islands. Fig. 4. Tle des Pins,
New Caledonia. All x20.
Plate 50. Gabrielona raunana goubini Robertson. Fig. 1. Out-
line of large shell, x17. Fig. 2. Apex, x33; p, protoconch. Fig
3. Aperture, x33; ar, apertural ridge; ps, palatal sulcus; uc,
umbilical channel.
[04-163]
54. Gabrielona
Operculum—See under G. raunana (species).
Types—The holotype (PI. 48, fig. 7) from Lifou,
Loyalty Islands, is in the Dautzenberg Collection
at the Institut royal des Sciences naturelles de
Belgique, Brussels. So also are 1,102 paratypes
from Lifou and Ile des Pins. Ten more paratypes
from Lifou are retained at the Academy of Nat-
ural Sciences of Philadelphia (no. 302624), and 3
from Lifou have long remained unidentified at
the United States National Museum _ (no.
422601), Washington, D. C.
Derivation of new name—Named for Goubin,
the collector who meticulously sorted out from
beach sand 1,085 of the shells.
Locality records (see map, Pl. 47)—. LOYALTY ISLANDS: Ile
Lifou (Goubin, IrSnB & ANSP; Moss, USNM). NEW CAL-
EDONIA: Ile des Pins (Lambert, IrSnB).
Robert Robertson
Phasianellidae
Gabrielona hadra (Woodring, 1928)
(Pls. 51-53)
Range—Middle Miocene or Plio-Pleistocene:
known only from the Bowden Formation, south-
eastern Jamaica, Greater Antilles. Presumably
was widespread in the Caribbean area. (On ap-
parent endemism in the Bowden Formation, see
W. P. Woodring, 1965, Science 148 (3672): 961-
963.)
Chief distinguishing characters—The shell at-
tained a larger size than that of any Recent spe-
cies in the genus. G. hadra is distinct also in hav-
ing a deeply embayed columellar lip below the
junction with the palatal wall. Otherwise, G.
hadra closely resembles G. sulcifera (the sculp-
ture of the second whorl is similar, the palatal
Plate 51. Gabrielona hadra (Woodring). Middle Miocene (or
Plio-Pleistocene), Bowden Formation, Jamaica. Fig. 4. Larg-
est known specimen of any Gabrielona (surface eroded). Figs.
5-7. Holotype. All x20.
[04-164]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
ul
on
Gabrielona
Plate 52. Gabrielona hadra (Woodring). Figs. 1-2. Outlines of
shells, showing variation, both x17. Fig. 2. Largest known
specimen of any Gabrielona (surface eroded). Figs. 3-4. Aper-
tures, showing variation; both x33; ar, apertural ridge; cdo,
lower (inner) columellar denticle; ps, palatal sulcus; uc, um-
bilical channel. Figs. 1 and 4. Holotype.
sulcus is near the middle of the aperture, and on-
ly the lower columellar denticle (edz) is present).
Additional differences from G. sulcifera: first
whorl averages smaller; spiral sculpture on sec-
ond whorl less prominent, disappearing before
the beginning of the third whorl; no spiral sulci
on later whorls; color patterns different.
Abundance—21 shells currently available, none
with opercula (36 paratypes not available; see
under Types).
Remarks—This is the only known fossil species
undoubtedly belonging in the genus, and clearly
is a precursor of the Recent Caribbean G. sulci-
fera. The relative abundance of G. hadra in the
Bowden Formation is noteworthy in view of the
rarity of its living descendant.
The surface of the shell is most resistant to cor-
rosion where there were white markings, which
on some shells are preserved as low projections.
Shell description—Attained length of 3.3 mm.,
width of 3.1 mm., and 4.6 whorls; spire angle
95°-120°; length invariably exceeds width (ex-
cept possibly for small shells), but outline vari-
able; fairly thick to thick, and opaque (fresh
shells might be translucent or transparent). Pro-
toconch slightly exsert, slightly inflated, smooth,
whitish, slightly demarcated from teleoconch,
1.1-1.2 whorls. First whorl 0.23-0.25 mm. in di-
ameter. Penultimate and last whorls: slight flat-
tening below suture and faint shoulder (PI. 52,
figs. 1-2). Sculpture of second whorl: 3 fairly low
spiral keels on first quadrant following proto-
conch, each about equal in prominence; keels
gradually becoming spiral cords, commonly with
2 more cords intercalated; very fine axial
threads; all sculpture gradually disappearing on
third or fourth quadrant, commonly with no spi-
ral sulci; on later whorls surface wholly smooth
except for fine axial growth lines. Surface shiny.
Colors: pale (faded?) reddish or purplish brown,
and white. Patterns: irregularly shaped whitish
subsutural patches (8-11 on last whorl), alternat-
ing with brownish areas; lower edges of patches
irregular, not correlated with position of palatal
sulcus; predominant below poorly demarcated
subsutural area: ground color pale reddish
brown; 2-3 spiral series irregular white markings
(commonly crescentic, concave towards outer lip)
that are smaller than the subsutural patches; 2 of
these series near (above and below) periphery
(upper one fairly uncommonly absent), and third
on base (6-9 markings); umbilical area com-
monly tinged with purple. Aperture shape like
[04-165]
56 Gabrielona
that of G. sulcifera; outer lip fairly thin to thick;
callus on upper parietal area thin to thick; pala-
tal denticle absent; apertural ridge (ar) shallow
to fairly deep in aperture (Pl. 52, fig. 3); pala-
tal sulcus near middle of aperture; upper colu-
mellar denticle (cd,) absent; lower columellar
denticle (cd,) present, prominent or low and
wide (Pl. 52, figs. 3-4). Columellar lip not thick-
ened, deeply embayed below junction with pala-
tal wall; fairly shallow slope into wide umbili-
cal channel; escarpment to left of umbilical
channel prominent, arising fairly high to high
off outer edge columellar lip, and commonly
the right-hand edge of a ridge; umbilicus wide.
Shell measurements (mm.)—
length width no. whorls
3.30 3.06 4.6 largest
2.01 2.44 4.2 average; holotype
2.10 2.04 4.0 fairly small
Synonymy—
1928 Tricolia (Eulithidium) hadra Woodring, Carnegie Instit.
Washington Publ. 385 (Miocene Mollusks from Bow-
den, Jamaica; Part I), pp. 16 [name listed], 420-421,
pl. 34, figs. 10-11.—1958, Robertson, Johnsonia 3(37):
253, 257 [provisionally referred to Gabrielona].
Types—The holotype of Tricolia (Eulithidium)
hadra Woodring (PI. 51, figs. 5-7), from near
Bowden, Jamaica, is at the United States Nation-
al Museum (no. 369556), Washington, D.C. This
presumably was one of the “37 specimens in the
Duerden Collection” mentioned by Woodring.
Plate 53. Geographical records of Gabrielona sulcifera Rob-
ertson (Recent) [round spots] and Gabrielona hadra
Robert Robertson
is
dl @ a ry a |
| ‘Sia sticrox " @ANTIGUA |
TAMAR EM Mio | ----ae eben ooo “pe |
eS aa | v \—'5
y |
= | =I a |
| , \s
CARIBBEAN | 2 \s
. SPD
Phasianellidae
The remaining 36, which can be considered para-
types, were not found in the paleontological col-
lection from Johns Hopkins University on deposit
at USNM (May, 1965). Twenty topotypes from
the Henderson collection are at USNM (no.
135509).
Fossil record (see map, Pl. 53, black triangle)— JAMAICA:
near Bowden, St. Thomas Parish (1894, J.B. Henderson, Jr.;
1899, J.E. Duerden; in thin bed imperfectly consolidated
gravel in a marly matrix; USNM).
Gabrielona sulcifera Robertson, new species
(Pls. 53-57)
Range—Recent: known only from off northwest-
ern Cuba, Greater Antilles, from the Virgin Is-
lands, and from Antigua, Lesser Antilles. Pre-
sumably widespread around high _ islands
throughout the West Indies, but very rarely col-
lected.
Chief distinguishing characters—This is the only
species in the genus having a shell with promin-
ent spiral sculpture. This begins on the second
whorl as keels, which soon become cords. On
later whorls the cords are reduced and_ broad-
ened to interspaces between sulci. These sulci
are absent at and near the periphery of the last
whorl of large shells. Also differs from other Re-
cent species in the genus as follows: very fine ax-
ial threads on second whorl; color pattern with
yt
be Boe oly
te
G. HADRA|™
OM. Mio.)
“a
(Woodring), its Middle Miocene (or Plio-Pleistocene) pre-
cursor [triangle].
[04-166]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
on
~l
Gabrielona
Plate 54. Gabrielona sulcifera Robertson. Antigua, Lesser
Antilles. Fig. 4. Holotype. All x20.
no demarcated subsutural area; palatal sulcus
near middle of aperture and not correlated with
color pattern on external surface; palatal denticle
can be present; only lower columellar denticle
(ed) present (abnormally, slight trace upper col-
umellar denticle); operculum with prominent spi-
ral cord at inner edge marginal spiral ridge.
For differences from G. hadra, see under that
species.
Relationships—Judging by the presence and
kind of sculpture (even though spiral and not
axial), the fairly thick, generally similarly-shaped
shells, and the two opercular resemblances, G.
sulcifera seems more closely related to G. rau-
nana than to the other two Recent species. How-
ever, the differences are far greater than those
between G. nepeanensis and G. pisinna.
Abundance—14 specimens available, only 1 live-
collected and with an operculum.
Habitat—The single live-collected specimen and
11 empty shells were all dredged together at an
unrecorded depth at Antigua, and were in “deep,
fine sand and shells.” To have been in or near
English Harbour, the depth must have been less
than about 20 fathoms (H.O. chart 366). The
empty, drilled, abnormal shell from 287 fathoms
off northwestern Cuba probably was adventi-
tious at that depth.
Shell description [see also section on ab-
normal shell|—Attains length of 2.4 mm., width
of 2.3 mm., and 4.3 whorls; spire angle 90°-
105°; length exceeds width, and even small
shells high-spired; outline fairly variable (Pl. 56,
figs. 1-3); fairly thick and slightly translucent to
slightly transparent. Protoconch slightly exsert,
slightly inflated, smooth, white, fairly prominent-
ly demarcated from teleoconch by slight varix,
1.1-1.2 whorls. First whorl 0.25-0.26 mm. in di-
ameter. Penultimate and last whorls: flattening
below suture and slight shoulder (PI. 56, figs.
1-3); commonly: slight spiral bulge around um-
bilical area (Pl. 56, figs. 4, 9). Sculpture of sec-
ond whorl: 3 spiral keels on first quadrant fol-
lowing protoconch varix, the middle keel (on
shoulder) the most prominent; keels less prom-
inent on second to fourth quadrant, gradually be-
coming spiral cords, with one or two more cords
intercalated; very fine axial threads (PI. 56, fig.
5). On later whorls, ‘cords are reduced and
broadened to interspaces between spiral sulci; 8-
12 sulci above suture on penultimate whorl. Last
whorl of small shells: 25-28 spiral sulci between
suture and base. Sulci absent at and near peri-
phery of large shells, and wholly absent near
outer lip where surface is smooth except for ax-
ial growth lines; sulci fairly regularly to irregu-
lary spaced (fairly uncommonly, in closely-spaced
pairs), especially variable below and near suture;
termination of sulci commonly abrupt. Surface
shiny. Colors: pale pinkish brown or orange-
brown, and white. Patterns: no demarcated sub-
sutural area; on middle whorls: irregular axial
bars at shoulder (0-10 per whorl), and near and
[04-167]
58 Gabrielona
above periphery (7-17 per whorl), developing
into wavy bands; on last whorl large shells: ax-
ially aligned, broadly wavy brownish bands aris-
ing from suture and extending to base, 6-11 on
last whorl, commonly coalesced into irregular
spiral areas on shoulder and on base, and com-
monly disjunct subperipherally; umbilical area:
white, with 5-9 (usually 6) axial or very steeply
descending brownish bands extending from mid-
dle of base; fairly rarely: all brownish marks very
pale and peripheral area uniformly whitish (PI.
54, fig. 4). Aperture slightly more rounded than
in other Recent species (less constricted near su-
ture); outer lip and callus on upper parietal
area fairly thick to thick; palatal denticle on most
large shells, commonly wide and faint; apertural
ridge (ar) shallow in aperture; palatal sulcus near
middle of aperture (lacking in smallest shell);
upper columellar denticle (cd;) absent; lower
columellar denticle (edz) present, commonly
wide (Pl. 56, figs. 4, 9). Columellar lip not
thickened, and steep slope into umbilical
channel but no escarpment; escarpment to left of
umbilical channel tairly prominent, arising fairly
high to high off outer edge columellar lip; broad
area to left of this escarpment, commonly with
several parallel threads; umbilicus fairly wide to
wide.
Plate 55. Gabrielona sulcifera Robertson. Off northwestern
Cuba. Abnormal shell. All x20.
Robert Robertson
Phasianellidae
Abnormal shell (Pl. 55)—The single empty shell
from 287 fathoms off northwestern Cuba (USNM
no. 94974) differs strikingly from all those from
Antigua, but seems to be conspecific. The proto-
conch and beginning of the second whorl are
normal, but the surface is increasingly corroded
as far as a growth line in the first quadrant of
the third whorl. Thereafter, the external surface
is smooth and shiny and lacks sulci. Another
growth line is at the beginning of the second
quadrant of the third whorl. I conclude from the
growth-lines, which are much more clear-cut
than on any other Gabrielona observed, that
growth of this shell was abnormal, perhaps be-
cause the animal lived in an unusual habitat
(depth?).
Other differences from the Antiguan specimens:
spire lower (spire angle ca. 115°); width exceeds
length (resembles G. pisinna in outline); thick;
coloration dark brick red and white; dark areas
large, with banding complex and greatly coal-
esced; apertural ridge (ar) fairly deep in aper-
ture; slight trace upper columellar denticle (ed,);
shallow slope from columellar lip into wide um-
bilical channel; no trace escarpment to left.
Shell measurements (mm. )—
length width no. whorls
2.42 2.27 4.3 largest (Antigua)
2.13 2:20 3.8 abnormal (N.W. Cuba)
1.98 1.93 4.0 average; holotype
1.33 1.17 She) smallest (St. Croix)
Operculum (Pl. 56, figs. 6-8)—The single oper-
culum available is different from all other known
Gabrielona opercula in having a prominent spiral
cord at the inner edge of the marginal spiral
ridge. The operculum of G. sulcifera is almost as
thick as those of G. raunana, and the opercular
whorl counts of these two species are also sim-
ilar. In outline, the operculum of G. sulcifera is
slightly more rounded than those of all the other
Recent species—a consequence of the more
rounded aperture. In the central region, a faint
spiral ridge separates the smooth outer spiral
area from the smooth central area. The callus is
thin, revealing some of the spiral suture at the
surface. The non-spiral columellar edge is steeply
beveled. The spiral edge seems to be corroded at
the external surface, and the sharp crest of the
spiral ridge may therefore be an erosional fea-
ture.
Radula (Pl. 57; 1 studied)—Attains length of 0.7
mm., width of 0.13 mm., and with as many as 30
[04-168]
May 30, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
Gabrielona — 59
fairly strongly curved transverse rows of teeth
(including a few nascent rows). Central monocus-
pid, with the small anterior portion of the base
in a cleft between the bases of the innermost
pair of laterals; the remainder of the base of the
central overlapping these (where they are juxta-
posed posteriorly). Five pairs of laterals, each of
these (like the central) with a small distal por-
tion that is monocuspid; the outer edge of
each base partly overlaps the base lateral to
it. As many as 34 pairs of marginals (lowest
count 26), the innermost large and massive
with lobed cusps, the distal portions gradually
becoming smaller and more slender outwards,
and with finer denticulations that finally are
absent altogether on the outermost teeth.
Synonymy—
1889 Phasianella (Eucosmia) brevis “Orbigny” Dall [Eucos-
mia brevis in plate caption] (not P. brevis Orbigny,
1842). Bull Mus. Comp. Zool. 18: 30 [listed], 351,
Plate 56. Gabrielona sulcifera Robertson. Figs. 1-3. Outlines
of large shells, showing variation, all x17. Fig. 1. Abnormal
shell from off northwestern Cuba. Fig. 3. Unusually high-
spired shell. Fig. 4. Holotype, showing position of sulci, x33.
Fig. 5. Apex, x60; p, protoconch. Figs. 6-8. Operculum of
pl. 19, fig. 10b [not North Carolinian specimens, which
are Tricolia thalassicola Robertson (deep water form)
and perhaps other species of Tricolia}.
1918 Eucosmia brevis (“Orbigny”)Cossmann (not P. brevis
Orbigny). Essais Paleoconch. Comp., Paris, livr. 11,
p 162, fig. 55 [outline copied from Dall, 1889].
1958 Gabrielona brevis (“Orbigny”) Robertson (not P. brevis
Orbigny, 1842). Johnsonia 3. (37): 257-260, pl. 138,
fig. 2 [radula], pl. 139, figs. 3-4 [operculum], pl. 140,
figs. 2-3, pl. 141 [not pl. 142, fig. 1].
Types—The holotype (PI. 54, fig. 4), the live-col-
lected specimen from Antigua, is at the United
States National Museum (no. 500636), Washing-
ton, D.C. Of the original 11 Antiguan paratypes,
7 are still at USNM (no. 659066), 2 are at the
Museum of Comparative Zoology (no. 188356), 1
is at the Academy of Natural Sciences of Phila-
delphia (no. 302625), and 1 (PI. 54, fig. 3) was
lost. The smallest paratype, fromm St. Croix, is
in Mr. Usticke’s private collection. The ab-
normal shell from Cuba is not a paratype.
Derivation of new name—Latin, sulcifer, bearing
furrows.
holotype. Internal surface, longitudinal section, and external
surface (respectively). All x33; c, lower edge of callus. Fig. 9.
Aperture, x33; ar, apertural ridge; ecd,, lower (inner) colu-
mellar denticle; pd, palatal denticle; ps, palatal sulcus; uc,
umbilical channel.
[04-169]
60 Gabrielona
Nomenclature—Following Dall (1889), who first
applied the name Phasianella brevis Orbigny,
1842, to the abnormal Cuban shell discussed and
retigured here (Pl. 55), I misapplied this name to
this species and transferred it to Gabrielona.
Subsequent study of the holotype (figured speci-
men) of P. brevis at the British Museum (Nat.
Hist.), no. 1854.10.4.282, has shown that this is a
depauperate Tricolia. This holotype, inadequately
illustrated in Robertson (1958, pl. 142, fig. 1),
will be treated in detail and refigured elsewhere.
Robert Robertson
Phasianellidae
Locality records (see map, Pl. 53, circular black spots)—
CUBA: off Bahia Honda, Pinar del Rio (23°2’N.; 83°13’W.;
287 fms.; Blake Sta. 21 [1877-78]; USNM). VIRGIN
ISLANDS: Christiansted Harbor, St. Croix (dredged 15 ft
[1 dead]; G. N. Usticke). LESSER ANTILLES: English
Harbour, Antigua (1918, J. B. Henderson, Jr., USNM, MCZ,
ANSP).
Erroneous locality record—Arenas de la Chor-
rera, Habana, Cuba (Robertson, 1958, pp. 259-
260, as G. brevis) [a juvenile Tricolia].
als. The cusps of the central and laterals are stippled. From
the holotype, Antigua, West Indies. x1500. Modified from
Robertson (1958, pl. 138, fig. 2, as Gabrielona brevis).
Plate 57. Gabrielona sulcifera Robertson. Half of one trans-
verse row of radular teeth, showing (from left to right) the
central between the innermost pair of laterals, the remaining
four laterals on the right side, and the entire row of margin-
[04-170]
May 30, 1973
EXCLUDED SPECIES
“Gabrielona” bruscasensis Weisbord, 1962
(PI. 58)
Range—Upper Miocene or Pliocene (or
younger?): known only from one locality in the
Playa Grande Formation (Maiquetia Member),
northern Venezuela.
Remarks—This species was described from a
single poorly preserved shell with a badly broken
columellar area. Even generic identification has
been a problem but the shell does seem to be a
phasianellid. However, it cannot be a Gabrielona
because the slight axial ridge revealing the posi-
tion of the withdrawn operculum is detectable,
and this is fairly deep in the aperture. I detected
no palatal sulcus or apertural denticles, and the
part remaining of the umbilical area indicates
that there was no Gabrielona-like umbilical
channel. The outer lip is more prosocline than in
any true Gabrielona. I conclude that G. brusca-
sensis probably is a young Tricolia.
The specimen lacks most of the characters dis-
tinguishing species of Tricolia. The apex is in
such poor condition that the whorls cannot be
counted accurately and the first whorl cannot be
measured. The shell is suffused with pale brown-
ish pink, but no color pattern is detectable. Sim-
ilar-sized shells of T. affinis cruenta Robertson
differ in outline (are more elongate, with a more
obtuse apex) and lack the umbilicus. G. brusca-
sensis is here considered a nomen dubium.
Synonymy—
1962 Gabrielona bruscasensis Weisbord, Bulls. American
Paleo., 42(193): 111, pl. 8, figs. 5-7 (Quebrada las
Bruscas [Playa Grande Formation], Distrito Feder-
al, Venezuela).
Type —The holotype is at the Paleontological
Research Institution (no. 26056), Ithaca, New
York.
Plate 58. “Gabrielona” bruscasensis Weisbord, probably a
young Tricolia. Upper Miocene or Pliocene [?], northern
Venezuela. Holotype. Both x20.
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
Gabrielona 61
“Gabrielona” sphaera Weisbord, 1962
(Pl. 59)
Range—Pliocene (or younger?): known only
from three nearby localities in the Mare Forma-
tion, northern Venezuela.
Remarks—This is a rissoacean. Although re-
sembling in outline a high-spired Gabrielona, it
cannot be one because the shell (for its size) is
thin, the periphery of the last whorl is lower than
in any known Gabrielona, the aperture is elon-
gate-ovate, there are no apertural denticles, and
neither is there a palatal sulcus. There is a wide
umbilicus and a broad umbilical channel that is
slightly angled at its lower left margin. The holo-
type has 3.4 whorls and the first whorl is 0.23
mm. in diameter. The callus on the upper parie-
tal area is incomplete medially. In coloration, the
shell is pale (faded?) amber and whitish near the
umbilicus and on the last part of the last whorl
where there are irregular amber markings. At the
apex, the beginning of the suture is tinged with
dark amber.
Synonymy—
1962 Gabrielona sphaera Weisbord, Bulls. American Paleo.,
42(193): 109-111, pl. 8, figs. 1-4 (near Quebrada Mare
Abajo [Mare Formation], Distrito Federal, Venezuela).
Types—The holotype is at the Paleontological
Research Institution (no. 26054), Ithaca, New
York. So also is the single paratype (no. 26055)
distinguished by Weisbord among eleven other
specimens identified with G. sphaera.
Plate 59. “Gabrielona” sphaera Weisbord, a rissoacean. Plio-
cene [?], northern Venezuela. Holotype. Both x20.
[04-177]
Published by
THE DELAWARE MUSEUM OF NATURAL HISTORY
Box 3937, Greenville, Delaware
19807, U.S.A.
[04-178]
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
June 1, 1973
{replacing vol. 2, no. 12, pp. 509-510; see stars]
Tectariinae 63
trations of these morphological differences in
Rosewater, 1970, and in this paper).
Most Tectarius s.s.and Echininus s.s. are living
today only in the East Indian area. With the
exception of Tectarius grandinatus whose range
extends eastward to Polynesia, all other species
are inhabitants of the raised, weathered coral reef
shorelines found in the Western Pacific Arc (per-
sonal observations, 1970). It can only be assumed
that this niche provides the requirements essen-
tial for the existence of these species:as they are to
be found nowhere else.
The fossil record provides very few clues to the
origin of these groups. There are only three Terti-
ary fossil species described from the Indo-Pacific
and these probably belong in three separate ge-
neric taxa. The oldest of these, T. songoense
Martin, from the Upper Eocene of Java, probably
represents nearly the earliest appearance of Tec-
tarius. AS mentioned in Part I, littorinid fossils
are exceedingly difficult to separate from Trochi-
dae and Turbinidae, and this is no less true for
Tectarius and Echininus. Probably, however,
these groups made their appearance in the early
Tertiary within the region where they have devel-
oped, and with the exceptions of Echininus (Tec-
tininus) nodulosus and Tectarius (Cenchritis)
muricatus, both of the western Atlantic, they have
remained there.
Opercula
The opercula of Indo-Pacific Littorinidae re-
quire special comment (see pl. 389). All are made
up of conchiolin, and those of members of the
subfamily Littorininae, including Littorina, its
subgenera, and Nodilittorina are usually pauci-
spiral and rather oval in shape with the nucleus at
the side and nearer one end (oligogyrous spiral
tvpe of Fretter, et al., 1962, pp. 79,80). In the
Echinininae, the basic plan of the operculum
differs from that of other littorines. It is the type
D
Plate 389. Opercula of Littorinidae and Trochidae.
Fig. A. Paucispiral operculum of Littorina (Littorinopsis)
scabra (Linne) from Mokuoloe Id., Kaneohe Bay, Oahu
(USNM 339388).
Fig. B. Mesospiral operculum of Tectarius rusticus (Philip-
pi) from Troughton Chain, northern Western Australia
(WAM 1787-69);
Fig. C. Multispiral operculum of Echininus cumingi (Phi-
lippi) from near Davao City, Mindanao, Philippines
(WAM 1566-70);
Fig. D. Multispiral operculum of Trochus niloticus Linne,
from Makuluva, Viti Levu, Fiji (USNM 531827).
Lines under each figure represent 5 mm.; stippled areas
are thickened and dark-brown in color; non-stippled areas
are light horn color and transparent.
of operculum which is called in other groups,
such as Trochidae, a miultispiral operculum
(polygyrous spiral type of Fretter, et al., ibid.)
although not so extreme as that figured by Fretter
(ibid., p. 80, fig. 43A; also see our pl. 389, fig. D.).
The operculum is circular in outline and moder-
ate to small in size. The nucleus is decidedly
central in location and growth proceeds outward
from the center in multiple, fairly evenly spaced
gyrations (pl. 389, fig. D). The operculum in Tec-
tarininae (fig. B) is intermediate in form between
Explanation to plate 388 (opposite page)
Figs. 1,2. Tectarius grandinatus (Gmelin) from Palmerston
Atoll, Cook Islands (USNM 685165)
Figs. 3,4. Tectarius tectumpersicum (Linné). Fig. 3, from
Stirling Isle, Treasury Ids., Solomon Islands (USNM
600370); Fig. 4, from “East Indies” (USNM 131450).
Figs. 5-7. Tectarius pagodus (Linne). Fig. 5, from “East
Indies” (USNM_ 18966); Fig. 6, from the Philippines
(USNM); Fig. 7, a young specimen from Polillo, Philip-
pines (USNM 311141).
Figs. 8.9. Tectarius rusticus (Philippi) from Buccaneer
Archipelago, Western Australia (USNM 684713).
Figs. 10,11 Tectarius coronatus Val. Fig. 10, from Pacific
(USNM 304587); Fig. 11, from Davao Bay, Mindanao,
Philippines (USNM 654034).
Figs. 12,13. Echininus cumingi cumingi (Philippi) from
Hervey Ids., Cook Ids. (USNM 42452).
Figs. 14,15. Echininus cumingi spinulosus (Philippi) from
Kadena Circle, Okinawa, Ryukyu Ids. (USNM 664658).
Figs. 16,17. Tectarius (Cenchritis) muricatus (Linné), from
Boca de Camarioca, Matanzas, Cuba (USNM 599944).
Figs. 18-20. Echininus (Tectininus) nodulosus (Pfeiffer).
Fig. 18, from Hog Island, Bahamas (USNM_ 603911);
Figs. 19,20, from Mujeres Harbor, Quintana Roo, Mexico
(USNM 662308).
(05-443a]
64 Joseph Rosewater
Littorininae and Echinininae, being large and
rather rounded in outline, with a slightly acentric
nucleus and having a number of gyrations more
than the paucispiral type but less in number than
the multispiral type. The Tectariine opercular
type is here termed the mesospiral or mesogyrous
spiral tvpe. The presence of the three opercular
types in Littorinidae may be considered to have
evolutionary significance, and possibly is related
to selection for a better aperture sealing mecha-
nism in animals which have considerable vertical
distribution on the shore line: in order, proceed-
ing from low toward higher shore habitats—
Littorininae, Tectariinae, Echinininae.
Reproduction
To my knowledge nothing is known concern-
ing reproduction in either Tectariinae or Echin-
ininae with the exception of Tectarius (Cen-
chritis) muricatus (Linné) which produces a pe-
lagic capsule (see Lebour, 1945, and Lewis, 1960,
references in Rosewater, 1970 p. 05-276). Field
and laboratory studies are needed to discover
details of the life histories of the remaining spe-
cies. However it is likely that most of these snails
also produce eggs encased in pelagic capsules
which undergo development in the sea. Abbott
(1954) noted that Lebour (ibid.) stated that some
of the Bermuda littorinids that live above high
tide line migrate to the water to spawn. It is
suspected that this also is the case with many of
the Indo-Pacific species.
Acknowledgments
The persons and institutions acknowledged in
Part I of this study (see Rosewater, Indo-Pacific
Mollusca, vol. 2, no. 11, p. 425) also are thanked
here. In addition, I acknowledge the following
for their help in making possible the examination
in the field of most of the species of Tectarius and
Echininus during the National Geographic Socie-
tv—Mariel King Memorial Expedition to the Mo-
luceas Islands, Indonesia, May to July 1970: the
late Mariel King, Mrs. Grace King, T. H. Richert,
C. Beal, C. M. Burgess, B. R. Wilson, and the
National Geographic Societv. The Government of
Indonesia graciously provided clearance for the
vessel Pele to work in the Moluccas Islands. Mr.
Kasim Moosa and Mr. Sukarno, both of the Insti-
tute for Marine Research, Djakarta, accompanied
the expedition and provided assistance of many
kinds.
Littorinidae
List of Recognized Taxa
Below is a list of the Tertiary fossil and Recent
species herein recognized as belonging in the
subfamilies Tectariinae and Echinininae. The
few fossil taxa are preceded by a dagger [#].
Family Littorinidae Gray, 1840
Subfamily Tectariinae, new subfamily
GENUS Tectarius Valenciennes, [1832]
Subgenus Tectarius Valenciennes, [1832]
coronatus Val., 1832. Type. Recent, wes-
tern Pacific
grandinatus (Gmelin, 1791). Recent, Pacific
islands
pagodus (Linné, 1758). Recent, western Pa-
cific
tectumpersicum (Linné, 1758). Recent, wes-
tern Pacific
rusticus (Philippi, 1846). Recent, northern
Australia
t songoense (K. Martin, 1931). Eocene, Java.
Subgenus
+Subditotectarius Ladd, 1966
trehderi Ladd, 1966. Type. Miocene, Mar-
shall Islands.
Subgenus Cenchritis von Martens, 1900
muricatus (Linne, 1758). Type. Recent,
tropical western Atlantic.
Subfamily Echinininae, new subfamily
GENUS Echininus Clench and Abbott, 1942
Subgenus Echininus Clench and Abbott, 1942
cumingi cumingi (Philippi, 1846). Type. Re-
cent, western Pacific
cumingi spinulosus (Philippi, 1847). Recent,
western Pacific
+ adelaidensis (Cotton, 1947). Pliocene, South
Australia.
Subgenus Tectininus Clench and Abbott, 1942
nodulosus (Pfeiffer, 1839). Type. Recent,
tropical western Atlantic.
[05-44-4a]
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
June 1, 1973
[replacing vol. 2, no. 12, pp. 511-512; see stars]
Tectariinae 65
Selected Bibliography
Abbott, R. T. 1954. Review of the Atlantic Periwinkles,
Nodilittorina, Echininus, and Tectarius. Proceedings of the
United States National Museum, vol. 103, no. 3328, pp.
449-464.
Argenville, A. J. D. d’. 1742. L’histoire naturelle —La Litho-
logie et la Conchyliologie —par—M. [A. J. D. d’ Argenville]
de la Société Royale des Sciences de Montpellier. Paris.
Bruguiere, M. 1792. Encyclopédie Méthodique, Paris, vol. 1,
p. 930.
Clench, W. J. and R. T. Abbott. 1942. The Genera Tectarius
and Echininus in the Western Atlantic. Johnsonia, vol. 1, no.
4, pp. 1-4.
Dance, S. P. 1967. Report on the Linnaean Shell Collection.
Proceedings of the Linnean Society of London, vol. 178, no.
1, pp. 1-24, 10 pls.
Dell, R. K. 1964. Marine Mollusca from Macquarie and Heard
Islands. Records of the Dominion Museum, vol. 4, no. 20,
pp. 267-301.
Deshayes, G. P. 1830. Encyclopédie Meéethodique, Paris, vol. 2,
p. 184.
Dodge, H. 1959. A Historical Review of the Mollusks. of
Linnaeus. Bulletin of the American Museum of Natural
History, vol. 118, Article 5, pp. 211-257.
Fretter, V., and A. Graham. 1962. British Prosobranch Mol-
luscs. Ray Society, London, xvi + 755 pp., 317 figs.
Habe, T. 1951. Littorinidae in Japan (1). Hlustrated Catalogue
of Japanese Shells, no. 14, pp. 87-93.
Habe, T. 1961. Coloured Ilustrations of the Shells of Japan
(II). Hoikusha Publishing Co., Ltd., Osaka, 183 pp., 66 pls.
Habe, T. 1964. Shells of the Western Pacific in Color, Vol. I.
Hoikusha Publishing Co., Ltd., Osaka. 233 pp., 66 pls
Kaicher, S. D. 1956. Indo-Pacific Sea Shells. Section 3. Lit-
torinacea, etc. Privately Printed, Washington, D.C., 8 pls.
and captions.
Keen, A. Myra. 1966. Tectarius (Mollusca: Gastropoda): Re-
quest for Validation in its Accustomed Sense. Z.N. (S.) 1754.
Bulletin of Zoological Nomenclature, vol. 23, part 4, pp.
179-180.
Kesteven, H. L. 1903. Notes on Prosobranchiata, No. II.
Littorinacea. Proceedings of the Linnean Society of New
South Wales, 1902, part 4, pp. 620-636.
Kira, T. 1959. Coloured Illustrations of the Shells of Japan.
Revised Edition, Hoikusha Publishing Co., Ltd., Osaka, ix
+ 239 pp.
Kira, T. 1962. Shells of the Western Pacific in Color. Hoikusha
Publishing Co., Ltd., Osaka, 224 pp.
Klein, J. T. 1753. Tentamen Methodi Ostracologicae sive
Dispositio Naturalis Cochlidum et Concharum, p. 25. Lug-
duni Batavorum.
Melville, R. V. and W. E. China. 1969. Opinion 871. Tectarius
Valenciennes, [1832] (Gastropoda): Validated Under the
Plenary Powers. The Bulletin of Zoological Nomenclature,
vol. 25, part 6, pp. 214-215.
Morch, O. A. L. 1852. Catalogus Conchyliorum Quae Reliquit
D. Alphonso D’Aguirra et Gadea Comes de Yoldi; fascicle 1,
p. 45.
Powell, A. W. B. 1951. Antarctic and Subantarctic Mollusca:
Pelecypoda and Gastropoda. Discovery Reports, vol. 26, pp.
47-196, text figs., 6 pls.
Prashad, B. 1925. Respiration of Gastropod Mollusks. Pro-
ceedings of the Twelfth Pacific Science Congress, pp.
126-143.
Rosewater, J. 1970. The Family Littorinidae in the Indo-
Pacific. Part I. The Subfamily Littorininae. Indo-Pacific
Mollusca, vol. 2, no. 11, pp. 417-506, 64 pls.
Rumphius, G. E. 1705. D’Amboinsche Rariteitkamer. Am-
sterdam, 340 pp., 60 pls.
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“Voyage”. Journal of Botany, June, pp. 1-4.
Troschel, F. H. 1856-1863. Das Gebiss der Schnecken, vol. 1,
Berlin, pp. vii + 252, pl. 1-20.
Watson, R. B. 1886. Report on the Scientific Results of the
H.M.S. Challenger, vol. 15, part 42, p. 576.
Wenz, W. 1938. Handbuch der Paliozoologie, vol. 6, part 1,
lfg. 3, pp. 241-480, figs. 472-1235.
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naturwissenschaftlichen Classe der Kaiserlichen Akademie
der Wissenschaften, Wien, I. Abth. vol. 80, pp. 496, 514.
[05-447a ]
66 Joseph Rosewater Littorinidae
Key to the Tectariinae and Echinininae
The following key is to the genera and sub- only, since Western Atlantic taxa are not treated in
genera of these two subfamilies. It is based upon detail in the present paper. For a key to the
the shell and externally observable characters. Littorininae, see vol. 2, no. 11, p. 430 [p. 05-278].
Page numbers are given for Indo-Pacific groups
LA OMe Wi DINER. 0 4 ox oot ered. eed 4ae 6S ERODES ee RRA ae aes 2
Lb: Sivell wet sa Bia ccs cs Gott ae ba eee eh We ee Rd Se Ge dew ed Gels ha ee dma ed 3
2a Shell spinose, with partially open spines, shell about as wide as high, operculum
pipe iS (see Pi SSO). oe es eka eo bea eh Oa eeds Homa dad Echininus p. 526
2b Shell not spinose, nodulose, higher than wide, operculum not multispiral . .. Cenchritis
3a Shell spinose, operculum mesospiral. Tectarius .......0.00 0000 ce ee ees p.o13
Sb Shell moculose,70S6il i oo ioc oo kaka he Pe eee dR KK RS Subditotectarius p. 524
3c Shell moderately spinose, operculum multispiral................005. Tectininus
ie
Plate 390. Type-species of Genera and Subgenera of Tectari- Miocene, Marshall Islands (Holotype, USNM 648342; 2.8
inae (Figs. 1-9) and Echinininae (Figs. 10-15) illustrating x 2.4 mm.).
sculpture, arrangement of spines, and presence or absence of Figs. 7-9. Tectarius (Cenchritis) muricatus (Linné);
umbilici. Matanzas, Cuba (USNM 599944, 26.1 x 17.9 mm.).
Figs. 10-12. Echininus (Echininus) cumingi (Philippi);
* Figs. 1-3. Tectarius (Tectarius) coronatus Val.; Davao Bay, Cook Islands (USNM 42452a; 17.2 x 17.5 mm.).
Mindanao, Philippines (USNM 654034; 23.1 x 17.8mm.). Figs. 13-15. Echininus (Tectininus) nodulosus (Pfeiffer);
Figs. 4-6. Tectarius (Subditotectarius) rehderi Ladd; early Cozumel Id., Mexico (USNM 662806; 14.4 x 12.2 mm.).
[05-448a ]
*
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
June 1, 1973
[replacing vol. 2, no. 12, pp. 513-514; see stars]
Tectarius 67
Subfamily Tectariinae, new subfamily
Genus Tectarius Valenciennes, [1832]
Type: Tectarius coronatus Val., 1832
The genus Tectarius sensu lato includes the
nominate subgenus, Tectarius, whose type-
species T. coronatus Val. (- T. papillosus ‘La-
marck’ of authors) has one of the more conserva-
tively sculptured shells of the group. Tectarius
sensu stricto is a wholly Indo-Pacific group, hav-
ing its present population center in the Western
Pacific Arc. The monotypic subgenus Cenchritis
von Martens contains only TJ. (C.) muricatus
(Linné), of the tropical western Atlantic (see pl.
388, figs. 16, 17). Subditotectarius Ladd, 1966, is
monotypic for the fossil T. (S.) rehderi Ladd, of
the Miocene of the Marshall Islands. Only spe-
cies belonging to Tectarius s.s. and Subditotec-
tarius will be considered here.
Tectarius appears more closely related to Lit-
torina than to Echininus because of greater simi-
larities in morphology, general shell characters,
the absence of a truly multispiral operculum, a
broader, less modified central radula tooth, and
the usual lack of an openly umbilicate shell.
Subgenus Tectarius sensu stricto
Moderately large, pyramidal to turbinate,
non-umbilicate littorinids with from rather
strongly spinose to nodulose or papillose shells;
generally living at or above high tide line. Radula
littorinoid, the central tooth somewhat narrowed,
the lateral tooth with an embayment and usually
developing a medial vertical ridge or partition. In
males the penis is large and well-supplied with
glands along most of its lateral edge, the remain-
der papillose, and with an open but deeply folded
seminal duct. Operculum rounded, mesospiral
(see Opercula in Introduction). Aperture plicate
within; with a columellar swelling or tooth.
Synonymy—
1798 Cidaris Réding, Museum Boltenianum, part 2, p. 84;
type-species by subsequent designation, Herrmann-
sen, 1847: Trochus pagodus Linné; non Cidaris
Leske, 1778, nor Swainson, 1840.
[1832] Tectarius Valenciennes, Coquilles, in Humboldt and
Bonpland. Voyage aux régions équinoxiales du
Noveau Continent, Observations de Zoologie, vol. 2,
p. 271; type-species by subsequent designation
Clench and Abbott, 1942: Trochus coronatus Valenci-
ennes.
1839 Pagodus Gray, in Molluscous Animals: The Zoology of
Captain Beechey’s Voyage, p. 141; type-species by
Monotypy and by absolute tautonymy, Monodonta
pagodus Lamarck [= Tectarius pagodus (Linné].
1840 Pagodella Swainson, A Treatise on Malacology, pp. 207,
219, 221 [refers to Pagodella echinata, nomen nudum]
351; refers to P. major Martini - Chemnitz, pl. 163,
figs 1541, 1542 [= Tectarius pagodus (Linné)] and to
T. tectumpersicum ibid., fig. 1543, 1544; type-species
here designated: Tectarius pagodus (Linneé).
1840 Echinella Swainson, ibid., pp. 207, 221, 352; refers to E.
granulata Swainson [nomen nudum] and to E. coro-
naria, Tableau Encyclopédique et Methodique, pl.
447, fig 6 [=Mondonta coronaria Lamarck = Tectar-
ius grandinatus (Gimelin)]; type-species by monotypy,
Tectarius grandinatus (Gmelin) [also see Clench and
Abbott, 1942]; not Echinella Bory St. Vincent, 1824.
1846 Fectaria Philippi, Abbildungen und Beschreibungen
Conchylien, Vol. 2, Litorina, p. 139; used in combina-
tion Fectaria pagodus, error for Tectarius Valencien-
nes:
1858 Hamus ‘Klein’ H. & A. Adams, The Genera of Recent
Mollusea, vol. 2, p. 656, refers to H. and A. Adams,
1854, vol. 1, p. 315; type-species here designated,
Hamus pagodus (Linne) [=Tectarius pagodus
(Linné)]; not Hamus ‘Klein’ R. B. Watson, 1886
{=Trochidae].
1899 Echinellopsis Rovereto, Atti della Societa Ligustica di
Scienze naturali e geografiche, vol. 10, p. 109; new
name for Echinella Swainson, 1840, not Bory St.
Vincent, 1524.
Nomenclature—Due to similarities between the
shells of Tectarius and some of the Trochidae,
there has been a tendency for some of the former
to be classified with the latter. This problem was
discussed by Keen (1966) who recommended that
the Zoological
Nomenclature validate Tectarius with the type-
Tectarius coronatus Valenciennes,
[1832], i.e., in its accustomed sense. Her petition
was granted in I.C.Z.N. Opinion 871 (Melville
and China, 1969). It was assumed in this Opinion
that the type-species of Tectarius, T. coronatus
International Commission. on
species
Valenciennes, is a synonym of T. grandinatus *
Gmelin. However, these are in actuality distinct
and geographically isolated species.
Another name which sometimes has been as-
sociated with Tectarius that has an exceedingly
long and complicated history is the genus Ha-
mus. It was mentioned originally by Klein (1753)
where its use was of course pre-linnaean. Bru-
guiéere (1792) gave a brief description, referring to
Klein, but listed no species. Deshaves (1830)
[05-449]
68 Joseph Rosewater
Littorinidae
declared it “a forgotten genus’, indicating that he
considered it unrecognizable. Mérch (1852)listed
it, this time in the synonymy of Littorina Férus-
sac, an invalid introduction (I.C.Z.N., Art. 11(d)).
The first valid use of Hamus was not until H. & A.
Adams (1858) used it as a senior synonym for
Tectarius (see synonymy). It was later used by
Wimmer (1880) and Watson (1886), the last being
a taxon of Trochidae. I have designated as type-
species of Hamus H. and A. Adams, 1858, H.
pagodus (Linné) and consider this genus to be an
absolute synonym of Tectarius Valenciennes.
The use of square brackets surrounding the
date for Tectarius Valenciennes, [1832] is reecom-
mended by the International Code of Zoological
Nomenclature in cases where the date of publica-
tion of a name has been determined on the basis
of external evidence (I.C.Z.N. Recommendation
92.A(3); also see Sherborn and Woodward, 1901;
Keen, 1966; and Opinion 871).
Tectarius coronatus Valenciennes, 1832
(PI. 388, figs. 10, 11)
Range—Philippines and Indonesia.
Remarks—The shells of well prepared and
cleaned specimens of Tectarius coronatus tendto
colorful for with the
orange-pink coloration of the last two whorls
contrasting with a purplish brown. subsutural
band. These colors do not show well in all speci-
mens, however, and are not very visible in un-
cleaned The
umbilicate common species of the southwest Pa-
be quite Littorinidae,
specimens. three large, non-
cific may be distinguished by the number of
major spiral rows of spines on the last whorl: 2 in
pagodus; 3 in tectumpersicum; and 4 in coronatus.
The closely-spaced stubby spines of coronatus also
separate it from the other two. These characteris-
tics do not of course help to distinguish it from T.
grandinatus, but other characters and the Polyne-
sian endemicity of the latter are helpful in this
case (see Remarks under grandinatus).
Habitat—Shore rocks and limestone cliffs 1-2
meters above high tide line (personal observa-
tions, Davao, Philippines, 1970).
Description—Shell reaching 39.7 mm (about
I'/2 inches) in length, broadly conical in shape,
average obesity about .76 (51 specimens range
from .68-.82);
heavily constructed, imperforate, and sculptured
mature specimens moderately
on most postnuclear whorls with four, fairly
closely-spaced rows of stubby, rounded, often
slightly upturned spines. External color generally
yellowish white on early whorls, becoming pink-
ish orange on penultimate and body whorls; area
of most posterior (subsutural) row of spines usual-
ly a contrasting purplish brown, and the same
dark color may appear in lines and dashes inside
outer lip of aperture; aperture tinted lighter pink-
ish orange. Base flattened, sculptured spirally
with nodulose cords, a larger separate row just
below periphery of body whorl. Whorls 6-8, flat-
sided excepting spines. Length of spire usually
greater than half the length of shell. Spire convex,
produced at an angle of from about 60-67°. Aper-
ture rounded-squarish; outer lip thickly produced
in mature individuals, strongly plicate within;
plicae not reaching edge of aperture; outer lip
tapering to a thin, crenulate edge; inner lip
smooth posteriorly, often stained a deeper orange
than rest of aperture, forming a tooth-like bulge
anteriorly, near junction with outer lip near base
of columella. Suture obscured by anteriormost
row of spines of preceding whorl. Primary sculp-
iural feature is the four spiral rows of spines.
Spines not particularly aligned axially, although
anteriormost 2 rows more so than others; from
17-23 spines per row on body whorl, bases of
anteriormost 3 rows of spines joined by low spiral
carinae. Posteriormost-but-one (3rd) row of
Plate 391. Tectarius coronatus
Figs. 1,2. Turbo rugosus Wood, lectotype, BM(NH)
1968370, 28.4 * 22.3 mm.
Figs. 3,4. Tectarius coronatus Valenciennes, Holotype,
MHNP (“Acapulco” [Luzon, Philippines]) 32.9 x 25.7
mam.
[05-450a]
*
INDO-PACIFIC MOLLUSCA, vol. 3, no. 14
June 1, 1973
[replacing vol. 2, no. 12, pp. 515-516; see stars]
Tectarius 69
spines protrudes farthest on spire whorls, but this
distinction largely lost on more mature whorls;
posteriormost (4th) row of spines obscured by
subsutucal purplish brown color band especially
on penultimate and body whorls. Secondary
spiral sculpture, between each row of spines, con-
sisting of raised cords, and overall spiral sculp-
ture of microscopic threads. Axial sculpture con-
sists of irregular flaky lines of growth. Operculum
moderate in size, circular, an average One measur-
ing about 7 mm. in diameter, thin, light-brown
with a dark-brown center, paucispiral, nucleus
about central. Periostracum not evident. Nuclear
whorls at least partially decollate in all specimens
examined, about 2, smooth, grayish white, first
postnuclear whorls showing early signs of spiral
striae and becoming nodulose. Radula littorinoid,
2-1-1-1-2; lateral tooth with a vertical partition
and an embayment typical of Littorinidae. Ani-
mal moderately large, littorinoid; penis large,
muscular and apparently highly extensible; semi-
nal groove in deep fold running along medial
edge to tip; tip vermiform, covered with papillae;
lateral edge of penis supplied with large number
of glands not extending onto vermiform tip. Re-
productive data and life history unknown.
Measurements (mm) (all Philippines)—
length width No. locality
whorls
39.7 27.6 7+ Cadao Id., Naro Bay, Masbate
35.4 25.2. 7+ Cadao Id., Naro Bay, Masbate
30.1 22.4 7+ San Miguel Bay, Ticao
95.9 D2, 8 Batag Id., Samar
23.9 16.3 7+ Bongao Channal, SW Sanga Sanga
Id., Sulu Archipelago
21.0 15.3 7+ Borongan, E side Samar
L725 130 7+ Papahag Id., Tawi Tawi Group
15.0 12.0 6+
13.5 10.8 7+
12.6 9.8 6+
Lee. 5.4 6+
5.4 6.8 5+
Borongan Village, E side Samar
Papahag Id., Tawi Tawi Group
Papahag Id., Tawi Tawi
Papahag Id., Tawi Tawi
Papahag Id., Tawi Tawi
Synonymy—
——— Monodonta papillosa of authors, not M. papillosa
Lamarck, 1822 [=Tectarius tectumpersicum (Linné,
1758) ].
1828 Trochus rugosus Wood, Supplement to the Index Tes-
taceologicus or a Catalogue of Shells, British and
Foreign, pl. 5, Trochus, fig. 7 (no locality given;
Mindanao, Philippines, here selected); lectotype in
BM(NH) 1968370, length 28.4 mm, width ( ca. ) 22.3;
not Litorina rugosa Menke, 1843 [= Nodilittorina aus-
tralis (Gray, 1926)]. Not Roding, 1798; Brown, 1815.
1832 Tectarius coronatus Valenciennes in Humboldt and
Bonpland, Voyage aux régions equinoxiales du
Noveau Continent, vol. 2, Coquilles, p. 271 (Acapulco
lin error] locality here corrected to Luzon, Philip-
pines); Holotype in MHNP.
1846 Litorina papillosa elegans Philippi, Abbildungen und
Beschreibungen Conchylien, vol. 2, p. 140, Litorina,
pl. 2, figs. 5, 7 (precise locality not given); figured
specimens from Cuming Collection BM(NH) [not
seen during 1968 visit] and Saul Collection, Cam-
bridge Museum.
1846 Litorina papillosa quadriseriata Philippi, ibid., p. 140,
Litorina, pl. 2, fig. 2 (Zanzibar [in error] locality here
corrected to Luzon, Philippines); type-specimen may
be in BM(NH) [not seen during 1968 visit]; refers to
“Trochus rugosus Wood Suppl. t.5, f. 7”.
Types—Although the name T. papillosus has
been applied to this species (see Kaicher, 1956)
the type-specimen of that species in the Geneva
Museum is unquestionably T. tectumpersicum
Linné, and papillosus is, therefore an absolute
synonym of tectumpersicum (q.v.). It also has
been referred to as Echinellopsis grandinatus
(Habe, 1961, p. 20; 1964, p. 28, both pl. 9, fig. 30)
which is an error of nomenclature for T. rugosus.
The first available name is Tectarius coronatus
Val. [1832], the holotype of which is in the
Paris Museum. The figured specimens of Philip-
pis elegans and quadriseriata may be in the BM
(NH) and/or the Cambridge Museum. They were
not discovered by me at the BMand may be lost.
Philippi’s figures are quite adequate for the
interpretation of the species and may be con-
sidered as representative of the lectotypes: ele-
gans, pl. 2, fig. 7; quadriseriata, pl. 2, fig. 2.
\Oo0° 120 140° 16¢
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Plate 392. Geographic distribution of Tectarius coronatus x
(Wood) in the Philippines and Indonesia.
[05-45 1a]
70 Joseph Rosewater
Records—PHILIPPINES: Port oe Mindoro; San
Miguel Bay, Ticao Id.; Cadao Id., Naro Bay, Masbate (all
USNM); Borongan Village ( (USNM, Del. Mus. ee ANSP,
MCZ): Babee ha both Samar (USNM); Samal Id., Davao Bay
(MCZ. USNM, WAM): Zamboanga, both Mindanao (ANSP,
Del. Mus. N.H.); Jolo Id. (MCZ); Tabawan Id. (ANSP);
Papahag Id. (USNM); Bongao Channel, SW Sanga Sanga Id.,
all Sulu Archipelago (ANSP). INDONESIA: Buka Buka Id..
Gulf of Tomini, Celebes (USNM).
Tectarius grandinatus (Gmelin, 1791)
(Pl. 388, figs. 1,2; pls. 393-395)
Range—The Cook, Society, Tuamotu and
Gambier Islands, southeastern Polynesia.
Remarks—Tectarius grandinatus, an inhabitant
of southeastern Polynesia, apparently is ge-
ographically isolated from the several other mem-
bers of the genus Tectarius in the Indo-Pacific,
the others being found in the East Indies. It is
quite distinct, morphologically, from pagodus
and tectumpersicum, but, interestingly, is very
¥& close in appearance to T. coronatus in general
matters of sculpture and external anatomy of the
animal. Superficially, grandinatus might be
thought more closely related to Cenchritis muri-
catus (L.) of the western Atlantic. However, the
radula, penial anatomy (Abbott, 1954) and a close
examination of shell sculpture and_ structure
causes me to reject that theory: grandinatus has
not been observed to be umbilicate, while muri-
catus sometimes is; the aperture of muricatus is
never thickened and plicate as it is in grandinat-
us; the operculum of grandinatus is rounded with
the nucleus near the center, that of muricatus is
more oval with the nucleus near the side.
The many similarities between coronatus and
grandinatus together with their spatial isolation
suggest the possibility that they may have
evolved from the same species stock. Although a
subspecific relationship may be indicated, the
two are here considered to have developed full
specific differentiation.
Habitat—Usually occurs on low islands within
the geographic range; on coral reef flats near the
high tide line, on jagged pieces of raised reef.
Description—Shell reaching 37.9 mm (about 1
1/9 inches) in length, elongate conical in shape,
average obesity about .72 (44 specimens range
from .62-.82); rather solidly and thickly con-
structed for its size, imperforate; mature speci-
mens with very deeply impressed suture and
rounded whorls; sculptured with four spiral rows
per whorl of stubby spines. External shell color
yellowish white; often coated with what appears
Littorinidae
to be a light-yellow to brown or rather dark
grayish brown periostracum which may wear thin
especially at tips of spines; no definite color
pattern apparent; occasional lines or splotches of
medium brown present; upper (most posterior)
part of aperture usually covered with a brown
glaze. Base moderately flattened, sculptured spi-
rally with nodulose cords. Whorls 7-9, moderately
rounded. Length of spire usually considerably
greater than half the length of shell. Spire convex,
produced at an angle of from 58-60°. Aperture
rounded; outer lip considerably thickened, pli-
cate within, tapering to a thinner, crenulate edge.
Inner lip with a thin, brown plage posteriorly,
forming a tooth-like bulge anteriorly, near junc-
tion with outer lip near base of columella. Suture
often deeply impressed, typically forming a chan-
nel between whorls. Primary sculptural feature is
the four spiral rows of spines. Spines not regu-
larly aligned axially, although anteriormost two
rows more-so than others; from about 17-25
spines per row on body whorl; bases of anteri-
ormost 3 rows of spines joined by low spiral
carinae; bases of posteriormost row of spines
usually separate; second from anterior row usual-
ly the smallest. Secondary spiral sculpture be-
Plate 393. Tectarius grandinatus (Gmelin, 1791).
Figs. 1,2. Trochus grandinatus Gmelin, lectotype in ZMC,
specimen figured by Chemnitz, Conchylien Cabinet, vol.
10, pl. 169, fig. 1639, from Palmerston Atoll, Cook Is-
lands, 32 x 22.4 mm.
Figs. 3-5. Monodonta coronaria Lamarck,
MHNG 1096/23, 41 x 27 mm.
holotype,
(05-452a]
May 30, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 14 71
List of Guide Tabs Cassidae 12-400
For *Tonnidae 13-400
Guide tabs are automatically sent to sub- Binder °Muricidae 13-800
scribers. Additional copies are available at the ° Vasidae 20-400
rate of 10 cents (U.S.) each. The present set of °Harpidae 20-600
16 costs $1.60 (U.S.).
Turridae 22-500
List of Issues 00-005 For Pinnidae 53-500
Eat Guide of Contents 00-100 Binder Tridacnidae 62-000
Render Phasianellidae 04-000 3 Index 90-000
l Littorinidae 05-250 Replaced Pages 95-000
Strombidae 09-650
Cypraeidae 10-400 °Text in preparation
List of Issues
Date of no. First page of
Volume 1 Subject Author Publication — pp. looseleaf
no. 1, pp. 1-14 Introduction Editors Mar. 31, 1959 14. = 00-003
15- 32. +Vasidae R. T. Abbott . 18 20-403
y, 33-146 Strombus R. T. Abbott Nov. 23, 1960 114 09-831
3; 147-174 — Lambis R. T. Abbott Sept. 28, 1961 38 10-051
4, 175-226 —-Pinnidae J. Rosewater . 52 53-501
2 227-346 Turrinae A. W. B. Powell Mar. 31, 1964 120 22-661
6, 347-396 _Tridacnidae J. Rosewater Apr. 30, 1965 50. 62-003
397-398 Strombus oldi W. K. Emerson . 2 09-881
399-400 — Strombus Listeri T. Okutani 2 10-001
401-402. Miocene Indian Strombus R. T. Abbott 2 09-945
403-406 Turrinae (replacement) A. W. B. Powell 4 22-947
407-408 — Issues and Changes Editors 2 00-005
GG 409-444 ‘Turrinae (concluded) A. W. B. Powell May 15, 1967 36 =. 23-101
445-454 Terebellum Jung and Abbott 7 1009-801
455-456 — Strombus wilsoni R. T. Abbott 2 09-919
8, 457-484 Cypraea (Zoila) Wilson and McComb Dec. 8, 1967 28 = 10-457
485-488 — Index to vol. 1 Editors ° 4 91-001
489-490 — Issues and Changes Editors 2 00-007
Volume 2
no. 9, l- 6 Title pages and Issues Editors Aug 30, 1968 6 00-009
7-202 Cassidae R. T. Abbott ° 196 = 12-409
10, 203-206 Masthead pages Editors Sept. 9, 1969 4 (00-003
207-416 Turriculinae A. W. B. Powell 7 210 23-191
i 417-506 — Littorininae J. Rosewater Nov. 30, 1970 90 = 05-261
12, 507-534 Tectariinae J. Rosewater Jan. 15, 1972 28 05-441
Volume 3
no.13, 1- 40 Drupa Emerson & Cernohorsky May 30, 1973 40 14-801
14 41- 62 Gabrielona R. Robertson 7 22 04-15]
63- 70 Tectarius (replacement) J. Rosewater June 1, 1973 8 05-443a
71- 74 Title page and Issues Editors 4 00-009a
[o0-009a]
INDO-PACIFIC
MOLLUSCA
Monographs of the Marine Mollusks of the World with Emphasis
on those of the Tropical Western Pacific and Indian Oceans
EDITED BY
R. TUCKER ABBOTT
VOLUME 3
Published by
DELAWARE MUSEUM OF NATURAL HISTORY
Box 3937, Greenville,
Delaware 19807, U.S.A.
74 Title Page Editors Volume 3
a
Notice
Looseleaf subscribers should place this title page in the first
binder immediately after the title page of volume 2. It will
be your record of the number of volumes in your looseleaf set.
[this sheet issued June 1, 1973, with vol. 3, no. 14]
7
November 27, 1973
5G OS
EH
Péfett
INDO-PACIFIC MOLLUSCA, vol.
3, no. 15 Patellidae
~~
our
LI
bes
SECTIONAL BRARY
DIVISION OF MOLLUSK
THE PATELLID LIMPETS OF THE WORLD
(PATELLIDAE)
by A. W. B. PowrELi
Auckland Institute and Museum
Auckland, New Zealand
Introduction
This monograph deals with the Patellidae, one
of the five families of limpets that comprise the
superfamily Patellacea. The patellid limpets are of
littoral and shallow-water occurrence and _al-
though widely distributed, cannot be considered
cosmopolitan, since they are absent from certain
extensive areas, namely, both coasts of North
America, the Caribbean, and South America,
north of Chile and Patagonia.
Although this work is concerned primarily with
the Indo-Pacific fauna, it is deemed necessary to
extend the scope to world coverage in order to
explain the otherwise apparently anomalous dis-
tributional patterns.
A complication is encountered with the decep-
tively similar shells of an allied family, the Ac-
maeidae, members of which are easily separable
from the Patellidae upon anatomical grounds, but
the shell of which usually has no character that
can be considered consistently diagnostic; hence,
with fossil limpets there is often an element of
doubt regarding family allocation.
Where patellids are absent, notably along the
North West American coast, the acmaeids take
over the corresponding littoral, ecological niche,
and in so doing, attain shell sizes very large for
acmaeids, which usually are of relatively smaller
size than patellids. A. striking instance of giant-
ism is the Californian Lottia gigantea Gray, 1834,
which may reach a length of four inches.
Limpets tend to vary greatly in size, shape,
sculpture and colour pattern, due to the ecolog-
ical factors involved, particularly the relative ex-
posure to wave stress and the nature of the sub-
stratum. Often, specific limits are apparent only
when extensive series from a number of stations
are studied.
Limpets featured frequently in early concholog-
ical works, but many of the species named are
difficult to determine with accuracy, since, for
the most part, they were based upon crude fig-
ures, inadequate descriptions, and with uncertain
locality data.
Under the heading of
cluded in the Patellidae”
“Species no longer in-
(pp.
84 to 87) 259
Plate 60. Gill structures in the Acmaeidae, Patellidae and
Lepetidae.
A=Acmaea_ virginea (Miiller); bp = branchial plume, be-
hind head.
B= Patella vulgata Linnaeus; ecc = gill cordon, complete.
= Cellana radians (Gmelin); gei gill cordon, interrupted
over head area.
D= Nacella (Patinigera) terroris
complete; epf= epipodial fringe
Patinigera ).
k= Lepeta coppingeri (E. A. Smith); no gills; respiration by
means of cilia-lined pallial groove.
(Fig. A from Fretter and Graham, 1962, p. 120, fig. 73. Fig. E
from Eales, 1923, p. 6, fig. 3).
(Filhol);
(only in
gece = gill cordon
Nacella and
[01-551]
e
es]
76 Patellidae
species, described as Patella are listed, and their
present familial location indicated. This list in-
cludes species now known to belong to the Ac-
maeidae, Lepetidae, Phenacolepatidae, Coccu-
linidae, _-Fissurellidae, Stomatellidae, Calyp-
traeidae, Capulidae, Hipponicidae, Muricidae,
Trimusculidae, Siphonariidae, | Umbraculidae,
Ancylidae, and even one considered to be based
upon one of the accessory plates of a member of
the Pholadidae.
Family Patellidae Rafinesque, 1815
The family Patellidae is one of three widespread
families——the Acmaeidae Carpenter, 1857, the
Patellidae Rafinesque, 1515, and the Lepetidae
Dall, 1869, all belonging to the superfamily
Patellacea.
Two other families, consisting of fossil species
only, are placed provisionally in the Patellacea;
they are the Metoptomatidae Wenz, 1935, of the
middle Silurian to middle Permian, and the Sym-
metrocapulidae Wenz, 1938, of the Triassic,
Jurassic and possibly Cretaceous.
A. W. B. Powell
The limpet shell is a simple shield or cap-
shaped structure, and is unfortunately a_shell-
form that is simulated by molluscs belonging to
several other gastropod orders. The one most
frequently mistaken for a patellid limpet is Si-
pPhonaria, a member of the air-breathing pul-
monates, being almost at the top rung of the
gastropod ladder of evolution, whereas the Patel-
lacea are located down towards the foot of the
ladder. In between are the fissurellids, crep-
idulids, capulids and umbraculids, all of which
have certain members that have limpetlike shells.
Limpetlike shells have developed independantly
in direct response to ecological necessity, being
the shell-form affording the greatest amount of
suction area for clinging to a rock surface, cou-
pled with a low profile to withstand wave stress.
Classification of the patellid limpets, therefore,
is dependent upon some knowledge of the animal.
Even the allocation of species to either the Ac-
maeidae or the Patellidae, the two major families
of the Patellacea, in many instances becomes
conjectural upon the evidence from the shell
alone.
Anatomical Outline of the Families and Major Genera of the Patellacea
Family Acmaeidae
A Leaf-shaped ctenidium only ...............
B Ctenidium present, plus gill cordon.
Jeans Setdnseceiiocenna asta ntoian acaba eae Acmaea
Radula; closely spaced pair of centrals alternating with wider
spaced pair of laterals; marginals vestigial or absent.
Gill cordon complete ...................
Gill cordon interrupted by head
Ey enh ot Saree SEEDER EIS ee cere te Scurria
sche tuicnsien date Se hain tahoe ae Oe ER: Lottia
Family Patellidae
C Ctenidium absent; replaced by gill cordon.
Radula; 4 or 5 central teeth, median one present, vestigial or
absent; lateral large, pluricuspid; marginals 3, weak or vestigial.
Gill cordon complete ..................
Radula; median central vestigial or absent; alternate pairs of long
curved centrals and laterals; marginals 3, weak to vestigial.
Gill cordon interrupted by head.
Epipodial fringe absent ............
Gill cordon complete.
Epipodial fringe present ..........
Family Lepetidae
D No gills; respiration by cilia-lined pallial groove.
Radula; large central, with prominent dentate cusp; no laterals;
2 functional Marcinals ciicsssoemsioriaes
Culiigoeeeaiunen ica mend easeenma siete Lepeta
November 27, 1973
Biology
The embryo of Patella hatches 24 hours after
fertilization. The trochophore is about 0.15 mm. in
diameter, with a tuft of erect apical cilia and two
rows of ciliated cells around the greatest perim-
eter of the larva. The cilia beat in clockwise man-
ner and rotate the top-shaped larva through the
water. Two days after fertilization the larva is
transformed into a pretorsional veliger, and both
shell and foot appear. Torsion then begins while
the larva is free-swimming. During the next stage
of about 30 hours the larva both swims and
crawls, and torsion is completed when the larva
is 3% to 4 days old, and this marks the end of its
pelagic life. The velum does not disappear until
the snail has been actively crawling for about the
third week. About this time the operculum is lost.
The shell of the veliger is a dextral coil of scarcely
one whorl, and this is soon replaced by a new
shell, after which the limpet’s post-larval life con-
tinues into the adult. (see Fretter and Graham,
1962, pp. 448-450 for a more detailed account).
The English Patella vulgata is a protandrous
hermaphrodite with most, if not all, individuals
starting life as male but later changing to female
at the age of one year or more. Investigations of
English populations of vulgata have shown that
90% of the limpets between 16 and 25 mm. in
length are male; in those about 40 mm. in length
the sexes are about equal; and in those 60 mm.
or more in length most are female (see Fretter
and Graham, 1962, p. 372).
Most patellids feed upon small species of living
algae, but some live upon giant kelp, where they
scrape away the surface tissue. The rock-dwelling
patellids feed with the head end moving method-
ically from side to side, while the radula operates
like a scythe. Patellids often travel up to four or
five feet in search of food, and usually manage to
return to their original resting places. This some-
times involves re-finding a site previously ex-
cavated in the rock that exactly fits the indenta-
tions of the shell margin. When rock faces have a
slight coating of silt, limpet journeys can be seen
quite plainly, and possibly the limpet uses its out-
ward track in finding its way home.
The age attained by limpets varies greatly
according to the species involved, the food poten-
tial of the particular habitat, and the zone in
which the species occurs. Fretter and Graham
(1962, p. 501), quoting Russell (1909), recorded
that Patella vulgata from certain established pop-
ulations in Scotland attained a length of about
29 mm. in the first year, during which time they
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patellidae = 77
reached sexual maturity. The estimated sizes
reached for each of the four succeeding years
were respectively 38, 44, 48 and 53 mm. On the
other hand, the same species under more favour-
able conditions at Plymouth reached a length of
53 mm. by the end of the second year. The esti-
mated life span for vulgata is about 15 years.
Very large and massive species, such as mexi-
cana and kermadecensis, probably live for a much
longer time, but size is governed also by the
availability of a suitable rock substrate, not un-
duly encroached upon by barnacles, corals, or
other encrusting marine growths. In the tropical
Indo-Pacific it is unusual to find large-sized lim-
pets associated with coral reefs, unless there are
intrusions of basaltic lava. In general large-sized
limpets are more frequent in cool-temperate wa-
ters than they are in the tropics.
Limpets that live in the upper tidal zone are
usually taller than individuals inhabiting the lower
tidal zone. Fretter and Graham (1962, p. 501),
quoting Orton (1932), correlated these differences
with the degree of exposure to desiccation. Lim-
pets living near low water mark remain exposed
for only a short period by the tide, but high-water
limpets are uncovered for the greater part of the
day, and this results in a general drying out of
the habitat.
To prevent desiccation the limpet must hold
firmly to the rock for the whole period that it is
out of water, and it is suggested that this con-
stant application of force by the attachment mus-
cles tends to pull in the mantle skirt, which is
responsible for new growth around the. shell
margin. This produces a taller and narrower shell
than that produced by an animal living lower
down in the tidal zone.
Limpets from very exposed situations often ap-
pear to be very different from examples living in
more sheltered situations. In the exposed ex-
amples, the shell becomes very flat in order to
withstand wave stress. As a result of this lowered
profile, the apex is located nearer to the anterior
end.
The colour patterns exhibited by juvenile lim-
pets tend to be less variable than those in more
mature examples of the same species, and often
provide more satisfactory diagnostic criteria. This
is especially evident in Cellana strigilis popula-.
tions from the southern islands of New Zealand
(Powell, 1955, pp. 65-67).
Surface erosion of the shell also greatly alters
the colour pattern. Thomson (1919, pp. 264-267)
described how erosion in Cellana radians often
eliminates the transverse “earlii” pattern, but
[01-553]
7S Patellidae
the pigmentation of the radial ribs survives due
to deeper impregnation of the shell substance.
With the shrinkage of the animal in senile ex-
amples, a thick, unicoloured callus is built up on
the inside of the shell, blotting out any macula-
tions that may have survived external erosion.
The Patellidae are the most successful and the
most advanced family of the Patellacea. The
change from a simple leaf-shaped ctenidium in
the Acmaeidae, to a gill cordon in the Patellidae
results in more efficient aeration. Although Scur-
ria and Lottia have developed a gill cordon they
still retain the acmaeid ctenidium. In the Patel-
lidae the ctenidium has entirely disappeared,
leaving the gill cordon as the sole means of res-
piration.
Patellid limpets have become so successful in
their chosen littoral environment that in some
locations, South Africa in particular, certain spe-
cies have become the dominant organisms of sev-
eral animal communities. In the “Cochlea zone”
of South Africa the species Patella cochlea is so
abundant that almost all other forms of animal
life are crowded out. A density of 1,300 individ-
uals of this limpet to the square yard has been
recorded, in so dense a concentration that as
many as 40 small individuals were found crowded
on top of a single large shell.
The radula
The radula in the Patellidae is long and narrow,
especially in Cellana, in which it sometimes has a
length of as much as four times that of the shell.
In situ it is concentrated in loose coils on the left
hand side when viewed from above. On the other
hand, Patella has a much shorter radula that folds
back upon itself at the nascent end.
The Patella radula consists of a strong or weak
or rarely absent median central, flanked by a pair
of centrals on either side, followed by a large
pRiricuspid lateral, and finally, three weak, slen-
der, functionless marginals. The latter may be
fused into a single plate. Well-developed cusps,
capped with a dark stain containing magnetite,
are present on the multiple centrals and the pluri-
cuspid laterals, but cusps on the remaining teeth
are small to vestigial, and colourless.
In Cellana and Nacella the radula differs from
that of Patella, in that the functional teeth are a
pair of long, large, centrals, closely-spaced, on
either side of a vestigial plate, which also may be
absent, alternating with a wider-spaced pair of
similar, well-developed laterals; the functionless
marginals are as in Patella.
A. W. B. Powell
In Patella the multiple centrals and the large
pluricuspid lateral have relatively short recurved
cusps, but in Cellana and Nacella, the alternating
pairs of centrals and laterals are very long and
project arcuately upward, almost at right angles
to the base. These long, strongly upcurved teeth
present difficulty in slide preparation, for they are
easily pressed at varying angles in mounting, and
thus may assume very different shapes. A satis-
factory solution to this problem is in the use of
cavity slides, that bridge the radula across, so
that the teeth assume their normal upright posi-
tion over the cavity, but are pressed sideways,
beyond the limits of the cavity, thus giving details
of denticles or indentations along the sides of the
teeth.
Many writers have endeavoured to use the
length of the radula in relation to the length of
the shell for separating three assumed closely
allied English Patella. The range of the means
arrived at by Fretter and Graham (1962, p. 495)
is tabulated below.
Plate 61. Fig. 1. Patella vulgata Linnaeus; England. Radula.
Fig. 2. Patella caerulea Linnaeus; Trieste. Radula, from
Thiele, in Troschel and Thiele, 1891, pl. 25, fig. 18.
[01-554]
November 27, 1973
vulgata intermedia aspera
Length of radula Sis
1.60-2.10
Length of shell
1.05-1.15
Unfortunately, in that work, there is no precise
indication of the actual identity of the species
termed Patella intermedia, there being four dif-
ferent usages of that combination by four differ-
ent authors.
Brian and Owen (1952, pp. 241-249) provided
a useful table, giving the valid name equivalents
for the nomenclature used in papers on European
Patellidae, published up until 1948 (see under
heading of Patella intermedia Auct.). Following is
the summary of Brian and Owen’s conclusions.
“Patella vulgata L. were collected from high-
and low-water levels on five different beaches
and the lengths of the shells and radula measured.
In all five localities the mean shell length was less
and the mean radula length greater at the higher
level, but the difference was not consistently sig-
nificant.”
“The complexity arose from the mergence of
two conflicting tendencies: one, intralevel, a posi-
tive regression of shell-length and radula. While
the former is no doubt a growth phenomenon, the
latter is probably an environmental effect—at
higher level exposure to desiccation and conse-
quently, prolonged adherence to the substrate,
caused a smaller shell base; reduced feeding time
caused a longer radula. These factors may have
resulted in confusion when comparing the values
of the radula fractions of species of Patella.”
Lowenstam (1962) has shown that the radular
teeth in the limpets, Acmaea, Lottia and Patella,
are capped with goethite, a dark opaque mineral
of high iron content. This gives the dark-stained
primary cusps of the patellacea a hardness of al-
most 5 on the Moho hardness scale, whereas the
radular hardness in the Littorinidae, the Fissurelli-
dae and certain trochoids is only between 2 and 3.
This hardness factor in tho radula enables lim-
pets to erode limestone and other rocks of com-
parable hardness, as well as the external surface
of other shells. Limpets frequently excavate deep
depressions in the rock to form a base of attach-
ment, and are known to return to their own par-
ticular site after a nightly foraging excursion.
Lowenstam (1962a) also suggested that since,
in the case of chitons, the dark stained denticle
caps show the presence of magnetite, then it is
possible that these magnetised teeth may serve as
a guidance system for the so-called homing in-
stinct of both chitons and limpets.
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patellidae 79
Geographical Distribution of the Patellacea
The family Patellidae has achieved a very wide
distribution, extending from near the Arctic Circle
to the Antarctic Circle, the western Atlantic, Med-
iterranean, West and South Africa, the Indo-
Pacific to as far north as northern Japan, east-
ward to the Hawaiian Chain and the Island of
Juan Fernandez, and southward to Australia, New
Zealand, the subantarctic of the southern end of
South America, the islands of the Southern
Ocean, and even a few locations adjacent to the
Antarctic Continent.
Strangely, there are no patellids on either the
east or the west coasts of North America, the
Caribbean and most of South America. The only
exceptions are the giant Patella (Ancistromesus)
mexicana from tropical Central America, obvi-
ously derived from the Indo-Pacific when the
former Tethys Sea was an open waterway around
the perimeter of the globe, and the cold water
Nacella-Patinigera complex that is strongly repre-
sented in southern Patagonia and Chile, and from
there spread eastward to the islands of the South-
ern Ocean.
The Recent distributional patterns for each of
the three families of the Patellacea are outlined
below (see plate 62).
Acmaeidae: The typical genus, Acmaea, is of
worldwide distribution, but has its greatest devel-
opment in western North America, where it takes
the place of the Patellidae, members of which are
absent from that area. Two other genera of the
Acmaeidae, Scurria and Lottia, are apparent fore-
runners of the Patellidae, since they have devel-
oped pallial gills, additional to the single gill-
plume of typical Acmaea. Their present range is
along the west coast of both North and South
America.
Patellidae: Typical Patella is well-represented
along the western coast of Europe, from the Lofo-
ten Islands to Great Britain, down to Spain, the
Mediterranean, the West African mainland and
off-shore islands, then southward to South Africa,
where typical Patella becomes more or less
merged into a composite local fauna of cold and
Wwarm-water subgenera. From there the genus
continues, in subgeneric form, across to the
warmer water island groups of the Indo-Pacific,
extending northward to Japan, eastward to the
west coast of tropical Central America, and south-
ward to the Kermadec Islands and the temperate
waters of southern Australia and Tasmania.
The genus Cellana is confined for the most part
to the warm and cool temperate waters of the
[01-555]
SO. Patellidae
A. W. B. Powell
Indo-Pacific, but in the New Zealand area it ac-
tually extends southward into subantarctic waters.
The farthest westward reached by Cellana is the
coast of Natal, the farthest eastward the island
of Juan Fernandez, off the coast of Chile, and the
farthest northward, Japan.
A third major genus in this family is Nacella,
containing the important subgenus Patinigera.
These are truly cold-water limpets, the greater
part of their range being subantarctic, but extend-
ing to the Antarctic by way of the Scotia Arc, and
also ranging northward up the Chilean coast to at
least Valparaiso, assisted in this by the upwelling
of cold water along that coast. The present con-
centration of the genus is in the Magellanic area,
and from there it spreads eastward, assisted in
this by the prevailing West Wind Drift. Many of
the species live upon the large kelps which pro-
vide an effective means of chance dispersal when
quantities of the weed are wrenched free and
drift before wind and current. The farthest east-
ward that this genus has established itself is Ker-
guelen Island, and for the subgenus Campbell
Island in the New Zealand southern islands.
The genus Nacella and its subgenus Patinigera
have a distinctive epipodial fringe, not found so
far in any other genus of the Patellacea. The
European Tertiary fossils attributed to Nacella
probably belong to other genera.
Lepetidae: This family consists of rather small
featureless white limpets, mainly from the deeper
waters of the Arctic Ocean, the north Atlantic,
the north Pacific, Mediterranean, Patagonia and
Antarctica. They are rather specialised, but not
necessarily highly advanced. There are no gills,
and respiration takes place through a cilia-lined
pallial groove. The presence of a large, broad-
based central tooth, with a conspicuous, broadly-
triangular, dentate cusp, no laterals, but a pair
of functional marginals, are radular characters not
found in the other two living families of the Patel-
lacea.
Fossil Occurrences of the Patellidae
Although a considerable number of fossil so-
called Patella species are encountered in liter-
ature, especially those from European Tertiary
localities, few of them can be assigned with cer-
tainty to that genus. The problem faced by
workers with Recent species of the Patellacea,
that of distinguishing between the Acmaeidae and
the Patellidae when the animal is unknown, is
even more a matter of conjecture when fossil
species are under consideration, especially with
those from the older formations, that have shell
features unlike those of living species.
The earliest species of the Patellidae that can
be generically identified with some degree of con-
Ft NORTHERN LIMIT of PATELLA
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140 1090
Plate 62. Geographical distribution of the Patellidae. Note
the almost entire absence of the family from North America,
and most of South America, with the exception of one species,
Ancistromesus mexicana, which occurs along the west coast
of Central America, and Nacella, with its subgenus Patini-
gera, in the southern part of South America, from where it
has drifted eastward over much of the Subantarctic and in
some areas of the Antarctic.
[01-556]
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patellidae $1
Plate 63. European and South Africa Patella
Figs. 1-3. Patella vulgata Linnaeus, 1758. Figs. 1, 2. Caldy
Island, South Wales. Fig. 3. [fracombe, England.
Fig. 4. Patella aspera Roding, 1798. Caldy Island, South
Wales.
Figs. 5-7. Patella caerulea’ Linnaeus, 1758. Isle of Capri,
Italy.
Fig. $8. Patella) caerulea Linnaeus, 1758 (forma crenata
Gmelin, 1791). Madeira.
[01-557]
Fig. 9. Patella (Cymbula) compressa Linnaeus, 1758. Kom-
metje, Cape Peninsula, South Africa. Lives on large kelp;
always sideways compressed,
Figs. 10, 11. Patella (Cymbula) miniata Born, 1778. Fig. 10.
False Bay, South Africa. Fig. 11. Port Elizabeth, South
Africa; beach shells bleach to bright pink.
Figs. 12-14. Patella (Olana) cochlear Born, 1778. Sea Point,
South Africa. Anterior end always constricted like a spout.
$2 Patellidae
A. W. B. Powell
fidence are Cellana carpentariana Skwarko, 1966,
from the late Neocomian lower Cretaceous of
Northern Territory, Australia, and Patella (Scutel-
lastra) cooperi (Powell, 1938) from the Otaian
Stage, lower Miocene of Motuihi Island, Auck-
land, New Zealand.
The above two records show at least that the
separation of Patella and Cellana is of long stand-
ing, and coupled with the radular divergence,
justifies Thiele’s division of the family into two
subfamilies, the Patellinae Rafinesque, 1815, and
the Nacellinae Thiele, 1929.
Doubtful species of Patellidae
Patella ? amuritica Wilckens, 1922
Range—New Zealand, Amuri Bluff, upper Cre-
taceous.
Remarks—This species is based upon a very
damaged and incomplete shell only 5 mm. in
length. The whole of the apical area is missing
and only a ring of shelly material remains. Even
Wilckens expressed doubt as to whether his gen-
eric determination was correct.
Synonymy—
1922 Patella ? amuritica Wilckens, N. Z. Geol. Surv. Pal. Bull.
no. 9, p. 5, pl. 1, fig. 8.
Patella guineensis Dunker, 1853
Remarks—The_ present writer has insufficient
West African material to evaluate Dunker’s spe-
cies, the name of which is preoccupied by Patella
guineensis Gmelin, 1791.
Synonymy—
1853 Patella guineensis Dunker, Ind. Moll. Guin. Infer., p. 49,
pl. 7, figs. 1-3; 19-21. Loanda, Guinea, West Africa
[Angola].
Cellana jutsoni Chapman and Crespin, 1934
Remarks—This species, from the lower Miocene
Plantagenet Beds of Albany, Western Australia,
is very doubtfully patellid. The presence of an
“obscure ridge, extending from the apex to the
posterior margin” suggests the fissurellid genus
Tugali or something akin to it.
Synonymy—
1934 “Cellana” jutsoni Chapman & Crespin, Journ. Roy. Soc.
West. Aust., vol. 20, p. 122, pl. 11, fig. 28.
Patella aspera Roding, 1798
(Pl. 63, fig. 4; pl. 68, figs. 3, 4)
Range—British Isles and Atlantic coast of
France.
Remarks—This “species” is not always readily
distinguished from vulgata. Typically it is more
elongated than vulgata, depressed, with the apex
nearer to the anterior end, and the primary ribs
are stronger and sharper, resulting in a more def-
initely corrugated margin. Other differences are
that the interior is porcellanous whitish, with the
head scar pale orange, radial colour lines are sub-
obsolete to obsolete, and the colour of the foot of
the animal is cream to orange, as opposed to
grey-green in vulgata.
Measurements (mm.)—
length width height
53.0 42.0 21.0 Caldy Id., South Wales
47.0 35.5 20.0 Caldy Id., South Wales
Synonymy—
1798 Patella aspera Roding, Mus. Bolten., vol. 2, p. 10 (refers
to Favanne, pl. 2, f. G).
1819 Patella aspera Lamarck, Anim. sans Vert., vol. 6, p. 327
(refers to Favanne, pl. 2, f. G).
1844 Patella athletica Bean, in Thorpe, Brit. Mar. Conch., p.
264, fig. 101).
1968 Patella aspera Lam., McMillan, Brit. Shells, Warne & Co.
Ltd., London. New York, p. 25, pl. 1, figs, 1, 3, 4.
Patella depressa Pennant, 1777
Range—South coast of England, Channel Islands
and Atlantic coast of France.
Remarks—This is the small, very depressed,
Siphonaria-like species, or form of vulgata, of
which Patella vulgata var. intermedia Jeffreys,
1865 is a synonym. Jeffreys described his variety
intermedia as “Shell rather smaller, flatter, and
oval, with finer ribs, and an orange crown; inside
golden-yellow or. tinged with flesh colour (oc-
casionally cream colour) in the centre, and beau-
tifully rayed toward the margin.” Forbes also re-
marked that the animal is black or dark-coloured.
Despite the detailed studies of both Fischer-
Piette and R. G. Evans, the taxonomic status of
both aspera and depressa in relation to vulgata is
still uncertain. Evans endeavoured to separate the
three as full species, upon minute differences in
the pluricuspid radula teeth, coupled with the
varying lengths of the radula for each. In dealing
with populations from the south of England Evans
admitted, that at the Isle of Wight, intermediate
forms were common but then remarked that to
the westward along the south coast three species
form discontinuous entities.
[01-558]
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patellidae 83
Synonymy—
1777 Patella depressa Pennant, Brit. Zool., vol. 4, p. 124, pl.
89, fig. 146.
1865 Patella, vulgata var. intermedia Jeffreys, Brit. Conch.
vol. 3, p. 237.
1923 Patella depressa Pennant, Tomlin, Journ. Conch., vol.
17, p. 34.
1935 Patella spp. Fischer-Piette, Systematique et biogeo-
graphie-Les Patelles d’Europe et d'Afrique du Nord,
Journ. Conchyl., vol. 79, pp. 5-66.
1952 Patella depressa Pennant, Evans, Proc. Zool. Soc., Lond.,
pp. 357-376.
Patella electrina Reeve, 1854
Remarks—This shell, described as coming from
Australia, is unlike any species known from that
area. Examination of the type specimen in the
British Museum (Natural History) revealed a
Patella of the caerulea series, very like the
Canary Islands Patella lowei d’Orbigny, so far as
one can judge from a single example.
Synonymy—
1854 Patella electrina Reeve, Conch. Iconica, pl. 22, fig.
55a, b.
Patella intermedia
Many writers have attempted to give taxonomic
status to the forms of both vulgata and caerulea
by providing detailed studies of the radulae, par-
ticularly its length in relation to that of the shell
(see also, under the heading “Radula,” in the in-
troduction), in small differences in the pluricuspid
laterals, and also in the colour of the foot of the
animal.
Unfortunately with many of these papers it is
difficult to correlate the results with the species
or forms investigated, due to a common fault of
many anatomists of placing little or no importance
upon the characters of the shell, and seldom illus-
trating the relevant shells.
Another source of confusion is in the nomen-
clature employed, specific names being frequently
cited without their respective authority and date.
A name that is quoted frequently, and around
which much useful data is associated is “inter-
media,” but which patellid of that name is in-
tended?—that of Knapp, 1557, Jeffreys, 1565, or
Bucquoy, Dautzenberg and Dollfus, 1852?
Brian and Owen (1952) endeavoured to rectify
this confusion by concluding that P. intermedia
Jeffreys, 1865, as used by Fischer-Piette (1935,
1938 and 1948), Eslick (1940) and Orton (1946)
was Patella depressa Pennant, 1777. They also
concluded that P. athletica Bean, 1844, as used
in the Plymouth Marine Fauna (1931) and by
Winckworth (1932) and Evans (1947) and P. de-
pressa, as used by Fischer-Piette (1935), Eslick
(1940) and Orton (1946) were all Patella aspera
Lamarck (now aspera Roding, 1798).
The earliest use of the name intermedia, in
association with Patella, is that of Knapp, in
Murray, 1857, proposed for a Channel Islands
shell, that appears to be a form of Patella vulgata
Linnaeus, 1758.
Patella lineata Lamarck, 1819
Remarks—This shell, from unknown locality,
was referred to Helcioniscus (now-Cellana) by
Pilsbry, but Delessert’s figures suggest a species
of the Acmaeidae. Pilsbry’s translation of Lam-
arck’s description follows:
Description—“Shell oval, convex, buff-brown,
painted with 10-12 yellow lines; excessively nu-
merous longitudinal close striae; vertex acute,
buff. Length exceeding one inch.” - 27 mm. (Mer-
mod).
Synonymy—
1819 Patella lineata Lamarck, Anim. sans vert., vol. 6 (1),
p. 331; Patella lineata Delessert, Rec. de Coq., pl. 23,
fig. 6.
1891 Helcioniscus lineatus Lamarck, Pilsbry, Man. of Conch.,
vol. 13, p. 153, pl. 73, figs. 85-87.
1950 Patella lineata Lamarck, Mermod, Revue Suisse de Zool.,
vol. 57, no. 34, p. 694 (remarks on the type).
? Patella nelsonensis Trechmann, 1918
Range—Nelson, New Zealand, lower conglom-
erates, lower slopes of range, upper end. of
Eighty-eight Valley, Kaihikuan Stage, middle Tri-
assic.
Remarks—The holotype and two paratypes are
in the New Zealand Geological Survey, Welling-
ton, but the preservation is not good enough to
show muscle scars, hence it cannot be deter-
mined, on the present material, whether the apex
is directed anteriorly or posteriorly. The species
remains a doubtful member of the Patellacea.
Synonymy—
1918 Patella (?) nelsonensis Trechmann, Quart. J. Geol.
Soc., vol. 73, pt. 3, p. 185, pl. 18, figs. S a, b.
1953 Patella nelsonensis Trechmann, Marwick, N. Z. Geol.
Surv. Pal. Bull. no. 21, p. 74, pl. 7, fig. 3.
Patella reussi K. Martin, 1879
Remarks—This species from the Miocene of Java
could be fissurellid. The figure shows a shell em-
bedded in matrix, and in consequence the interior
[01-559]
$4. Patellidae
of the shell, with its muscle impressions, is un-
known.
Synonymy—
1879 Patella reussi K. Martin, Die Tert. auf Java, Leiden,
p. 87, pl. 12, fig. 9.
Patella spectabilis Dunker, 1853
Remarks—The_ description and_ figures of
Dunker’s species suggest a form of Patella lowei
d’Orbigny, 1839. However, Dunker’s name is pre-
occupied by Patella spectabilis Gmelin, 1791.
Synonymy—
1853 Patella spectabilis Dunker, Ind. Moll. Guin. Infer., p. 39,
pl. 6, figs. 7-9. Loanda, Guinea, West Africa [Angola].
Species no longer included in the Patellidae
The following species, originally referred to the
Patellidae, are now known to belong to other
families. This list includes only species that the
writer has been able to evaluate, either per-
sonally, or upon the authority of subsequent
revisers. There remains a considerable number of
patellid names yet to investigate, but since many
of these have been inadequately described, sel-
dom figured, and often without locality data, it is
probable that most, eventually, will have to be
considered indeterminate.
Species prefixed by an asterisk (*) are the sub-
ject of a note at the conclusion of this section.
The writer has compiled a manuscript list of
over 1,250 patellid names encountered in the
preparation of the present work, but it is withheld
from publication at this stage, since it cannot be
considered reasonably complete, without a more
thorough appraisal of European species, the fossil
ones in particular. A dagger (+) precedes fossil
species in this list.
achates Reeve, 1855, Patella ............ccccccccseeeeeseeneeeeee Acmaeidae
acinaces Lea, 1846, Patella .........0...ccc cece eee ceeeeeeee ees ? Pholadidae
aculeata Gmelin, 1791, Patella 2.0.0.0... ccsseseens Caly ptraeidae
adunca Perry, 1811, Patella... eee Fissurellidae
aenigmatica Mabille, 1895, Patella 00.0000... Acmaeidae
aeruginosa Middendorff, 1847, Patella (Acmaea) ... Acmaeidae
afra Gmelin, 1791,: Patellas wsisssccsiuscsctnaantessaccrantvens Trimusculidae
albescens Anton, 1839, Patella ...........cccccceeceeeeeeeeees Acmaeidae
albicosta C. B. Adams, 1855, Patella ..............:008 Acmaeidae
albicostata Reeve, 1855, Patella .....0.....0..cccccceeeeee ees Acmaeidae
alticostata Angas, 1865, Patella w.........ccccccceeecseeee tees Acmaeidae
alveus ‘Conrad, 183i... Patel adits nnconaaiisaiennecenanats: Acmaeidae
ambigua Wood, ISIS, Patella ...........0.ceeeeeeeeeees Fissurellidae
amoena, Say; 1822, Patella siiisisscissssccanctecsensedesasesdess Acmaeidae
ancyloides E. Forbes, 1840, Patella oo... Lepetidae
ancyloides Middendorff, 1847, Patella (Acmaea) ... Acmaeidae
angulata Wood, 1828, Patella ............cccceeeeeeeee Calyptraeidae
angusta Gmelin, 1791, Patella oo... cece Fissurellidae
A. W. B. Powell
antillarum Philippi, 1849, Patella (Acmaea) ............ Acmaeidae
antiquata Linnaeus, 1767, Patella ...................0 Hipponicidae
apertura Montagu, 1803, Patella .......00000.0.....008 Fissurellidae
araneosa Gould, 1848, Patella ......00.ccccceceeeeeeeeeeeees Acmaeidae
araneosa Reeve, 1855, Patella .......0.0..cceceeceee cece eee Acmaeidae
araucana d’Orbigny, 1841, Patella .....000...00...c0 Acmaeidae
asmi Middendorff, 1847, Patella ......0.0...0.00ccccccee Acmaeidae
atricapilla Dillwyn, 1817, Patella oo... Fissurellidae
auricula Gmelin, 1791, Patella 0.0.00... Calyptraeidae
auricula W. Wood, 1828, Patella. .........0.....:c0cee Stomatellidae
australis Lamarck, 1819, Patella ...0......ceees Hipponicidae
avellana Gmelin, 1791, Patella .........0000...cccccceeeee Fissurellidae
axiaerata Verco, 1912, Patella ............ccccccsscsssseeeeeees Acmaeidae
balanoides Reeve, 1855, Patella .....0...000.cccceeee Acmaeidae
barbadensis Gmelin, 1791, Patella ....000.....0c.ccce Fissurellidae
biradiata Reeve, 1855, Patella ........0.00ceccceeeeeeeees Acmaeidae
borneensis Reeve, 1855, Patella ..........00....ccccceceeeeee Acmaeidae
borniana Helbling, 1779, Patella .................0000cceeeees Acmaeidae
caeca Muller, 1776, Patella ........0...cccccccecceeseeeeesseees Lepetidae
calamus Crosse & Fischer, 1864, Patella ............00... Acmaeidae
callosa Hombron & Jacquinot, 1841, Patella .......... Acmaeidae
campaniformis Blainville, 1825, Patella ............ ? Siphonariidae
campbelli Filhol, 1880, Patella ....00...00...cccee Acmaeidae
cancellata Gmelin, 1791, Patella ......00.00.ccceee Acmaeidae
candida Couthouy, 1838, Patella 0.0.0.0. Lepetidae
cantharus Reeve, 1855, Patella ........cccccccccceeeeeeeceeeeee Acmaeidae
cassida Dillwyn, 1817, Patella oo... Hipponicidae
casta Carpenter, 1866, Nacella .............cccccceecceeeeees Acmaeidae
ceciliana d’Orbigny, 1841, Patella oo... Acmaeidae
cerea Mller, 1842, Patella .......0..cccccccccssesseeeeneeees Lepetidae
chilensis Blainville, 1825, Patella .....0..00..000. ? Siphonariidae
chinensis Linnaeus, 1758, Patella ......000...00..0..... Caly ptraeidae
cimeliata Reeve, 1855, Patella ....0..0.0000000 eee Acmaeidae
cinis Reeve, 1854, Patella .........0.0.ccccccccessecseseseeeeeeeees Acmaeidae
cinnamomea Gould, 1846, Patella ................ Phenacolepadidae
clealandi J. Sowerby, 1822, Patella .....0....cce Acmaeidae
clypeus T. Brown, 1827, Patella .......00.c.cccccceeeeeeeee Acmaeidae
cochleata Dillwyn: 1807, oc.ccccssssieag cose esnintessscaecess Capulidae
coffea Reeve, 1855, Patella ............c0cccccccessssseeeeeeeeees Acmaeidae
compressiuscula Karsten, 1849, Patella ..........0....... Acmaeidae
concentrica Middendorff, 1851, Patella ..............000.. Lepetidae
concepsionis Lesson, 1831, Patella ............. Acmaeidae
concinna Lischke, 1870, Patella ..................0000ce cee Acmaeidae
conica Defrance, 1825, Patella ...............cccccceeeeeceeees Acmaeidae
conica Gould, 1846, Patella ........00cccccccceeeeeeeee Acmaeidae
conulus Dunker, 1882, Patella .........cccccccccceeecceceeeeeee Acmaeidae
corrugata Reeve, 1854, Patella ...........ccccccecceece sence Acmaeidae
craniolaris Roding, 1798, Patella oo... eee Fissurellidae
crebrestriata Verco, 1904, Nacella .....0...000ccccceeeee Acmaeidae
crepidula Linnaeus, 1764, Patella .....0..00..00.006. Calyptraeidae
cruciata Linnaeus, 1758, Patella ....0.0.000000.cceeee Acmaeidae
crystallina W. Wood, 1828, Patella ...............0000 Fissurellidae
cubensis Reeve, 1855, Patella ..........c0cccccccecseeeeeeeees Acmaeidae
cumingii Reeve, 1854, Patella ..........0..0..ee Acmaeidae
diaphana Reeve, 1854, Patella .........c0cccecccsseceees Acmaeidae
dichotoma Anton, 1839, Patella ......0000....cccceeeeeeeees Acmaeidae
digitale RGding, 1798, Patella .......0.0ccccccecceeeeeeeeees Acmaeidae
discors Philippi, 1849, Patella (Acmaea) ................ Acmaeidae
elegans Philippi, 1849, Patella ....0..0000.0cccee Acmaeidae
emarginuloides Philippi, 1868, Patella .........00...0.0..... Lepetidae
equestris Linnaeus, 1758, Patella... Calyptraeidae
exilis Philippi, 1849, Patella ......0..00. 0c ccecceeceeeee Acmaeidae
fenestrata Reeve, 1855, Patella ......0000.cccccccceeeeereee Acmaeidae
fimbriata Gould, 1846, Patella ...0.00000....cccceeceeeeeeee Acmaeidae
fissura Linnaeus, 1758, Patella .............0.000cc cece Fissurellidae
fissurata Dillwyn, 1817, Patella 0.0.0... Fissurellidae
fissurella O. F. Miiller, 1776, Patella ........0.0c0ccccce. Fissurellidae
floccata Reeve, 1855, Patella .......00ccccccccceeeeeeeeeeeees Acmaeidae
fluviatilis Gmelin, 1791, Patella ......0.0ccccccceceeeeeee Ancylidae
forbesii J. Smith, 1839, Patella oo... cece Lepetidae
fornicata Linnaeus, 1758, Patella oo... Calyptraeidae
fulva O. F. Miiller, 1776, Patella ...0.0..0ccccceeeee Lepetidae
fungoides Réding, 1798, Patella v.00 Acmaeidae
[01-560]
November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae 85
Plate 64. South African Patella
Figs. 1-3. Patella (Patellona) granatina Linnaeus, 1758. Sea Figs. 7-9. Patella (Patellona) oculus Born, 1778. Buffel’s
Point, South Africa. Bay, Cape Peninsula, South Africa.
Figs. 4-6. Patella (Patellidea) granularis Linnaeus, 1758. Figs. 10-11. Patella concolor Krauss, 1848. Near Durban,
Sea Point, South Africa. Natal.
[01-561]
S86 Patellidae
A. W. B. Powell
galathea Lamarck, 1819, Patella ........00........ Phenacolepadidae
goreensis Gmelin, 1791, Patella oo... Calyptraeidae
graeca Linnaeus, 1758, Patella ...........000.....cccceeeeeees Fissurellidae
grammia Philippi, 1847, Patella... Acmaeidae
granostriata Schrenck, 1867, Patella .........0.....0000 Acmaeidae
granulata Philippi, 1848, Patella... Acmaeidae
grisea Gmelin, 1791, Patella... Siphonariidae
grisea Réding, 1798, Patella ...........0..00cccceeeceeceeeeeees Fissurellidae
haliotoidea Réding, 1798, Patella ...........0....6. Calyptraeidae
hepatica Pritchard & Gatliff, 1903, Patella ............. Acmaeidae
heptagona Blainville, 1825, Patella .........0.......08 Acmaeidae
heroldi Dunker, 1882, Patella .................ccccccesceeeeees Acmaeidae
hiantula W. Wood, 1828, Patella ....0.............00ccce Fissurellidae
°hochstetteri K. Martin, 1879, Patella) .................. Fissurellidae
incisa Dillwyn, 1817, Patella 20.0000... Fissurellidae
inconspicua Gray, 1843, Patella ..........0.. Acmaeidae
indica W. Wood, 1828, Patella .................ccceee Umbraculidae
inradiata Reeve, 1855, Patella ..............00...ccceeeeeeeee ? Acmaeidae
insessa Hinds, 1842, Patella 0.0.0... eee eee Acmaeidae
insignis Menke, 1843, Patella ...........0.......cccceccecee eee Acmaeidae
instabilis Gould, 1846, Patella ..........0000000.ccc cece Acmaeidae
intertexta ROding, 1798, Patella ......000.00..cc eens Acmaeidae
jacksoniensis Reeve, 1855, Patella ......0.....cceeee Acmaeidae
jamaicensis Gmelin, 1791, Patella p. 3715 w........... Acmaeidae
jamaicensis Gmelin, 1791, Patella, p. 3730 ............ Fissurellidae
javanica Lamarck, 1819, Patella ................c28. Siphonariidae
kochi Philippi, 1549, Patella oo... eee Acmaeidae
laciniata Reeve, 1855, Patella ........0......0ccccceeee eee Acmaeidae
laciniosa Linnaeus, 1758, Patella ..............ce Siphonariidae
lacunosa Reeve, 1855, Patella ................000ccccee eects Acmaeidae
lacustris Linnaeus, 1758, Patella ............0..cc eee Ancylidae
lamanonii Schrenck, 1867, Patella ..................000:0005 Acmaeidae
lanx Reeve, 1855, Patella ............cccccccccccsssseeeeeeeeeeeees Acmaeidae
laqueare W. Wood, 1828, Patella ...........:ce ? Acmaeidae
latistrigata Angas, 1865, Patella 0... ee Acmaeidae
lentiginosa Reeve, 1855, Patella ..............::c0ccceeeeeeeees Acmaeidae
lepas Gmelin, 1791, Patella ............0.05. Muricidae (Thaidinae)
leucophaea Philippi, 1849, Patella .......000....... Acmaeidae
leucopleura Gmelin, 1791, Patella ................0.00 Acmaeidae
leucopleura Reeve, 1855, Patella .........0.. cee Acmaeidae
lima Reeve, 1855, Patella .............00cccceececeeeeeeeee seen es Acmaeidae
limbata Réding, 1798, Patella ...........ceeees Fissurellidae
lineata Philippi, 1849, Patella... Acmaeidae
*luchuana Pilsbry, 1901, Patella... Acmaeidae
luctuosa Hombron & Jacquinot, 1841, Patella ........ Acmaeidae
lutea Linnaeus, 1758, Patella ................eee Stomatellidae
macroschisma Lightfoot, 1786, Patella .................. Fissurellidae
macroschisma Dillwyn, 1817, Patella ..........0....0.... Fissurellidae
mamillata (Nuttall) Reeve, 1855, Patella ................ Acmaeidae
mauritiana Pilsbry, 1891, Helcioniscus .................5 Acmaeidae
melanoleuca Gmelin, 1791, Patella .....................:008 Acmaeidae
melanoleuca Reeve, 1855, Patella 0.0.0.0... Acmaeidae
merceauxi Deshayes, 1861, Patella ..............:ee Acmaeidae
minima Gmelin, 1791, Patella ......0..........c0cccceeeee ees ? Acmaeidae
mitella R6ding, 1798, Patella .................cccceeeeeeeeeees Fissurellidae
mixta Reeve, 1855, Patella ............ccceeeeeeeeeeeeeeeeeeees Acmaeidae
miilleri Dunker, 1875, Patella (Tectura) ................. Acmaeidae
muricata Brocchi, 1814, Patella .................e. Calyptraeidae
mytiliformis Gmelin, 1791, Patella oo... Calyptraeidae
navicula Reeve, 1854, Patella oo... ccc eeeees Acmaeidae
neptuni Dillwyn, 1817, Patella 00... Calyptraeidae
°nigrosulcata Reeve, 1855, Patella RE Rem Acmaeidae
nimbosa Linnaeus, 1758, Patella .................:0 Fissurellidae
noachina Linnaeus, 1771. Patella .....0..0...............0- Fissurellidae
nodosa Born, 1778, Patella ...........60ccccceceeeeeeeeeeeeeeees Fissurellidae
notata Linnaeus, 1758, Patella .....0.......0..ccceeeeeeeeeee Fissurellidae
nubecula Linnaeus, 1758, Patella .....................000 Fissurellidae
nummularis R6ding, 1798, Patella ..... .... Fissurellidae
nuttalliana Reeve, 1855, Patella ...............ceceeeeeee Acmaeidae
obliquata Koenen, 1892, Patella (Acmaea) ....... ? Cocculinidae
obscura Hombron & Jackquinot, 1541, Patella ...... Acmaeidae
occidentalis Reeve, 1855, Patella ...........c.ccccceeeceeeees Acmaeidae
octoradiata Gmelin, 1791, Patella Fissurellidae
onychites Menke, 1843, Patella ..........0...0cccccceeeseees Acmaeidae
topea Reeve, 1854, Patella... eeeeeeeees Acmaeidae
oregona Nuttall, 1839, Patella .......0000.00ee .. Acmaeidae
pallescens Philippi, 1849, Patella (Acmaea) ............ Acmaeidae
tpallida Gould, 1859, Patella ....0...cccceeeee ... Acmaeidae
papillaris Réding, 1798, Patella .............00.ccceeee Acmaeidae
parasitica d’Orbigny, 1841, Patella ............. . Acmaeidae
parva da Costa, 1778, Patella .........0.cccceee ... Acmaeidae
patina Eschscholtz, 1847, Patella (Acmaea) ............ Acmaeidae
pectinata Linnaeus, 1758, Patella .........000..0..00.. Siphonariidae
pediculus Philippi, 1846, Patella .......00.000.00.cccce Acmaeidae
peltoides Carpenter, 1864, Nacella. .... .... Siphonariidae
penicillata Reeve, 1855, Patella ........0..0.ceeee Acmaeidae
perforata Gmelin, 1791, Patella .........0.c eee Fissurellidae
persona Eschscholtz, 1847, Patella (Acmaea) ......... Acmaeidae
personata T. Martyn, 1788, Patella (non binom.) Fissurellidae
perversa Gmelin, 1791, Patella... Umbraculidae
personoides Middendorff, 1849, Patella (Acmaea) Acmaeidae
peziza W. Wood, 1828, Patella ............ ees Calyptraeidae
phyrozonias Gmelin, 1791, Patella oo... Fissurellidae
picta Gmelin, 1791, Patella .......ccceeeeeeereees Fissurellidae
pileolus Dillwyn, 1817, Patella wo... cee Fissurellidae
pileolus Middendorff, 1849, Patella (Acmaea) ....... Acmaeidae
plana Philippi, 1849, Patella 00... eee Acmaeidae
plana Reeve, 1855, Patella ..............0cccccccceceeeeeeeeeeens Acmaeidae
poculum W. Wood, 1828, Patella ..............0:00 Calyptraeidae
pretrei d’Orbigny, 1841, Patella ....00...00.cc eee Acmaeidae
tprofunda Deshayes, 1863, Patella .............0.:c Acmaeidae
punctatissima Philippi, 1849, Patella... Acmaeidae
punctulata Gmelin, 1791, Patella ................ccccceeeeees Acmaeidae
puncturata Lamarck, 1819, Patella 0.0... Acmaeidae
pustula Gmelin, 1791, Patella .........00.0000c ccc Fissurellidae
pustulata Helbling, 1779, Patella 0... eee Acmaeidae
pygmaea Dunker, 1882, Patella .............00000cc eee Acmaeidae
rosea Gmelin, 1791, Patella ............000ccc cece ceeeeeeeee eee Fissurellidae
rosea Dall, 1872, Nacella ? wc... ccccccecccceesceeeesceeneeeees Acmaeidae
roseoradiata Verco, 1912, Nacella crebrestriata ..... Acmaeidae
rubella O. Fabricius, 1780, Patella .........000......cceeee Acmaeidae
rubeola R6ding, 1798, Patella ........000000.....0ceeeee Fissurellidae
rudis R6ding, 1798, Patella ...............0ccccceseeeeseeeeeees Fissurellidae
rugosa Réding, 1798, Patella 0.0.0.0... Capulidae
saccharina Linnaeus, 1758, Patella... Acmaeidae
scabra Gould, 1846, Patella (Lottia) ......000000.0....... Acmaeidae
scabra Reeve, 1855, Patella ..........eceeeeeeseeeeeeeeeeees Acmaeidae
scapula Martyn, 1789, Patella (non binom.) ............ Aplysiidae
schrenckii Lischke, 1868, Patella ........00......00:cee Acmaeidae
scurra Lesson, 1831, Patella ...........cccccceeeeeeeeeeees Acmaeidae
scutellata W. Wood, 1828, Patella oo... Calyptraeidae
senilis R6ding, 1798, Patella... Umbraculidae
sinensis Gmelin, 1791, Patella... ee Calyptraeidae
sinica Gmelin, 1791, Patella ........0000..ccececeeece neces Umbraculidae
sinuosa Brocchi, 1814, Patella... Capulidae
solandri Colenso, 1844, Patella .............0cccceececeeeeeeeee Acmaeidae
spectrum Reeve, 1855, Patella ............c cc ceeceeeeeceeees Acmaeidae
spinosa Gmelin, 1791, Patella .............. cece eeeeeeeeeee Fissurellidae
squamulata Renier, 1804, Patella ........... . Calyptraeidae
stella Lesson, 1831, Patella .......00.000ccccccceccceeeeeeeees Acmaeidae
stellaris Reeve, 1855, Patella .......00...cccccccececcceeeeeee ees Acmaeidae
stipulata Reeve, 1855, Patella ...............cccccccseeeeeeee Acmaeidae
stowae Verco, 1906, Nacella ..........cc cece ..... Aemaeidae
striata Reeve, 1855, Patella 0.0... eters Acmaeidae
sturnus Hombron & Jacquinot, 1841, Patella .......... Acmaeidae
subspiralis Carpenter, 1864, Nacella ..........0.00. Siphonariidae
sulcata Borson, 1520, Patella ..... . Hipponicidae
tectum Dillwyn, 1817, Patella
tectumchinensis R6ding, 1798, Patella
. Calyptraeidae
. Calyptraeidae
tenera C. B. Adams, 1845, Patella ......00000000000.... Acmaeidae
tenuicostata Michelin, 1838, Patella ........000000000.0.... Acmaeidae
tessellata O. F. Miiller, 1779, Patella .........0000000000... Acmaeidae
testudinalis O. F. Miiller, 1776, Patella .......00.00.. Acmaeidae
textilis Gould, 1846, Patella (Lottia) ........0000..000000. Acmaeidae
tranquebarica Gmelin, 1791, Patella ........0.0.. Acmaeidae
triangularis Carpenter, 1866, Nacella paleacea ....... Acmaeidae
[01-562]
November 27, 1973
tricarinata Linnaeus, 1767, Patella .........00...0.0.. Hipponicidae
tricostata Gmelin, 1791, Patella oo... Hipponicidae
trochiformis Gmelin, 1791, Patella ..................... Calyptraeidae
trochoides Dillwyn, 1817, Patella .........00...0... Calyptraeidae
tuberculifera Lamarck, 1819, Patella ................ ? Siphonariidae
turcica Réding, 1798, Patella... Calyptraeidae
umbellata Gmelin, 1791, Patella ...........00........... Umbraculidae
umbellata della Chiaje, 1830, Patella ................. Umbraculidae
umbonata Reeve, 1855, Patella .......00.....00...c00cece cece Acmaeidae
uncinata Reeve, 1855, Patella ....................cccccecceeeees Acmaeidae
undulata Réding, 1798, Patella oo... ee Caly ptraeidae
ungarica Linnaeus, 1758, Patella .......0........ceeee Capulidae
unguis Linnaeus, 1758, Patella ........0.0.....000....000eces. Fissurellidae
unguis J. Sowerby, 1816, Patella... eee Capulidae
unguisalmae Lesson, 1831, Patella .....00..000...0...000...08. Acmaeidae
verriculata Reeve, 1855, Patella ....000....00000.cccecceeeeeee Acmaeidae
vespertina Reeve, 1855, Patella ..........cceeeeeceeees Acmaeidae
victoriae Gatliff & Gabriel, 1922, Patella ................ Acmaeidae
victoriana Singleton, 1937, Patella ....0...00...cc eee. Acmaeidae
virginea O. F. Muller, 1776, Patella .....00....000000...... Acmaeidae
viridula Lamarck, 1819, Patella ..........00.......cc0cceees Acmaeidae
zebrina Lesson, 1831, Patella o.....0.0...000cccccccceeeeeeeeee Acmaeidae
Notes relevant to the above non-patellid species
Patella hochstetteri K. Martin, 1879
Remarks—This species, from the Miocene of
Java, is a Hemitoma, family Fissurellidae; in fact
its author likened it to Hemitoma notata (Lin-
naeus, 1755).
Synonymy—
1879 Patella hochstetteri kK. Martin, Die tert. auf Java, Leiden,
p. 86, pl. 12, fig. 10.
Patella luchuana Pisbry, 1901
Remarks—Habe determined that the radula and
gill structure of this Ryukyu Islands limpet prove
it to belong to the Acmaeidae.
Synonymy—
1901 Patella luchuana Pilsbry, Proce. Acad. Nat. Sci. Phila.,
vol. 53, p. 202.
1957 Collisella luchuana Pilsbry, Habe, Proc. Malac. Soc.
Lond., vol. 32, p. 207.
Patella nigrosulcata Reeve, 1855
Remarks—This shell, described from unknown
locality, has since been identified as a Western
Australian Patelloida, family Acmaeidae, and thus
has nothing to do with Patella (Scutellastra) stel-
laeformis, where it was assigned as a variety by
Pilsbry, 1891. The species is usually found at-
tached to the backs of large Patella (Scutellastra)
laticostata Blainville.
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patellidae 87
Synonymy—
1855 Patella nigro-sulcata| Reeve, Conch. Iconica. pl. 30,
figs. 84 a, b.
IS91 Patella (Scutellastra) stellaeformis var. nigrosulcata
Reeve, Pilsbry, Man. of Conch., vol. 13. p. 100, pl. 66,
figs. 66, 67.
1955 Patelloida nigrosulcata Reeve. Macpherson, Proc. Royal
Society of Victoria, vol. 67 (2), p. 241.
Patella opea Reeve, 1854
Remarks—Dr. Myra Keen has shown that the
originally cited locality for this species, “Sand-
wich Islands” - Hawaii, is incorrect, and that the
species is a synonym of the West American
Acmaea fascicularis Menke, 1851.
Synonymy—
1854 Patella opea Reeve, Conch. Iconica, pl. 29, figs. 79 a, b.
1958 Acmaea fascicularis Menke, Keen, Sea Shells of Tropical
West America, p. 244.
Patella pallida Gould, 1859
Remarks—This Recent Japanese species is now
recognised as belonging to the Acmaeidae.
Synonymy—
1859 Patella pallida Gould, Proc. Boston Soc. Nat. Hist., vol.
Ge 162:
1952 Tectura pallida Gould, Kuroda & Habe, Check List Rec.
Mar. Moll. Japan, p. 89.
Patella profunda Deshayes, 1863
Remarks—This Recent Reunion Island species is
now known to belong to the Acmaeidae.
Synonymy—
1863 Patella profunda Deshayes, Moll. Réunion, p. 44, pl. 6,
figs. 15, 16.
1942 Patelloida profunda Deshayes, Tomlin & Stephenson,
Proc. Malac. Soc., London, vol. 25, p. 6.
Helcioniscus profundus
var. mauritiana Pilsbry, 1891
Remarks—This Recent species and its variety
from the island of Mauritius belong to the Ac-
maeidae.
Synonymy—
IS9L Helcioniscus profundus var. mauritiana Pilsbry, 1891,
Manual of Conchology, vol. 13, p. 150, pl. 65, figs
97-99.
[01-563]
SS Patellidae
A. W. B. Powell
List of Recognized Taxa
SUBFAMILY Patellinae
Genus Patella Linnaeus, 1758
Subgenus Patella Linnaeus, 1758
vulgata Linnaeus, 1758. Type. Europe
aspera Roding, 1798. Europe
depressa Pennant, 1777. Europe
ferruginea Gmelin, 1791. Mediterranean
baudonii Drouet, 1858. Azores
rustica Linnaeus, 1758. S. Europe-Mediter-
ranean
piperata Gould, 1846. Madeira and Cape Verde
Ids.
caerulea Linnaeus, 1758. Mediterranean, Por-
tugal, Azores and Canary Ids.
moreleti Drouet, 1858. Azores
lowei dOrbigny, 1839. Canary Ids.
gomesti Drouet, 1858. Azores
Subgenus (not known)
candei d'Orbigny, 1839. Canary Ids.
citrullus Gould, 1846. Madeira
concolor Krauss, 1848. Natal and eastern South
Africa
depsta Reeve, 1855. St. Paul and Amsterdam
Ids.
rangiana Rochebrune, 1882. Cape Verde Ids.
tkaffraria Rennie, 1930. Cretaceous, South
Africa
Subgenus Patellona Thiele, in Troschel & Thiele,
1891
granatina Linnaeus, 1758. Type. South Africa
oculus Born, 1778. South Africa
adansonii Dunker, 1853. West Africa
canescens Gmelin, 1791. St. Helena
lugubris Gmelin, 1791. West Africa and Cape
Verde Ids.
plumbea Lamarck, 1819. West Africa
safiana Lamarck, 1819. Algeria to West Africa
Subgenus Patellidea Thiele, in Troschel & Thiele,
1891
granularis Linnaeus, 1758. Type. South Africa
Subgenus Cymbula H. & A. Adams, 1854
compressa Linnaeus, 1758. Type. South Africa
miniata Born, 1778. South Africa
Subgenus Olana H. & A. Adams, 1854
cochlear Born, 1778. Type. South Africa
Subgenus Scutellastra H. & A. Adams, 1854
argenvillei Krauss, 1848. South Africa
barbara Linnaeus, 1758. Type. South Africa
longicosta Lamarck, 1819. South Africa
tabularis Krauss, 1848. South Africa
exusta Keeve, 1854. Mauritius
subsp. pica Reeve, 1854. Mauritius to Sey-
chelles
flexuosa Quoy & Gaimard, 1834. Indo-Pacific,
Andamans to Tuamotus
subsp. optima Pilsbry, 1927. Japan
kermadecensis Pilsbry, 1894. Kermadec Ids.
taurorae Fleming, 1973. Middle Oligocene, New
Zealand
tucopiana (Powell, 1925). Tikopia, Melanesia
laticostata Blainville, 1825. south West Australia
peronii Blainville, 1825. southern Australia
chapmani Tenison Woods, 1875. South Austra-
lia to New South Wales
thamiltonensis (Chapman
Lower Pliocene, Australia
tcooperi (Powell, 1938). Lower Miocene, New
Zealand
Subgenus Ancistromesus Dall, 1871
mexicana Broderip & Sowerby, 1829. Type.
West Mexico
t fuenzalidai Herm, 1969. Pliocene, Chile
& Gabriel, 1923.
Genus Helcion Montfort, 1810
Subgenus Helcion Montfort, 1810
pectunculus (Gmelin, 1791). Type. South Africa
Subgenus Ansates Sowerby, 1839
pellucidus (Linnaeus, 1758). Type. Western
Europe
P tella (Bergh, 1871). Sargasso Sea
Subgenus Patinastra Thiele, in Troschel & Thiele,
1891
pruinosus (Krauss, 1848). Type. South Africa
dunkeri (Krauss, 1848). South Africa
SUBFAMILY Nacellinae
Genus Cellana H. Adams, 1869
eucosmia (Pilsbry, 1891). Red Sea
radiata (Born, 1778). India to Philippines
subsp. capensis (Gmelin, 1791). Natal to Zan-
zibar
subsp. enneagona (Reeve, 1854). Madagascar
to Japan
subsp. orientalis (Pilsbry, 1891). Indonesia;
Japan; Marquesas
tdeformis (K. Martin, 1883). Miocene, Java
karachiensis (Winckworth, 1930). Gulf of Oman
to Karachi
livescens (Reeve, 1855). Type. Mauritius
pricei Powell, new species. Samoa and New
Hebrides
garconi (Deshayes, 1863). Reunion and Mada-
gascar
[01-564]
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patellidae 89
Plate 65. Patella of the subgenus Scutellastra
Figs. 1-3. Patella) (Scutellastra) barbara Linnaeus, 1758.
Figs. 1, 3. Buluga Bay, East London, South Africa. Fig. 2.
Port Alfred, South Africa.
Fig. 4. Patella (Scutellastra) argenvillei: Krauss, 1548. Sea
Point, South Africa.
Figs. 5-7. Patella (Scutellastra) longicosta Lamarck, 1519.
Kommetje, Cape Peninsula, South Africa.
Figs. $-9. Patella (Scutellastra) flexuosa Quoy & Gaimard,
1834. Fig. 8. Paea, Tahiti. Fig. 9. Wake Island.
Fig. 10. Patella (Scutellastra) flexuosa subspecies optima
Pilsbry, 1927. Waki, Satsuma, Japan; young example.
Figs. 11-13. Patella (Scutellastra) peronii Blainville, 1825.
Fig. 11. Swansea, Tasmania. Figs. 12-13. Shellharbour,
New South Wales, Australia.
[01-565]
90 - Patellidae
(Cellana cont'd)
testudinaria (Linnaeus, 1758). Andaman Ids. to
New Caledonia
vitiensis Powell, new name. Fiji
grata (Gould, 1859). Japan and Korea
mazatlandica (Sowerby, 1839). Japan and Ry-
ukyu Ids.
nigrolineata (Reeve, 1854). Japan
toreuma (Reeve, 1855). Japan to Philippines
exarata (Reeve, 1854). Hawaiian Ids.
talcosa (Gould, 1846). Hawaiian Ids.
tahitensis (Pease, 1868). Tahiti and Pitcairn
ardosiaea (Hombron & Jacquinot, 1841). Juan
Fernandez Id.
conciliata Iredale, 1940. Queensland
turbator Iredale, 1940. South Queensland
tramoserica (Holten, 1802). South Queensland
to South Australia
solida (Blainville, 1825). Tasmania to South
Australia
tcarpentariana Skwarko, 1966. Lower Creta-
ceous, North Australia
tcudmorei Chapman & Gabriel, 1923. Lower Mi-
ocene, Victoria
thentyi Chapman & Gabriel, 1923. Lower Plio-
cene, Victoria
analogia Iredale, 1940. Lord Howe Id.
howensis Iredale, 1940. Lord Howe Id.
craticulata (Suter, 1905). Kermadec Ids.
denticulata (Martyn, 1784). New Zealand
flava (Hutton, 1873). New Zealand
ornata (Dillwyn, 1817). New Zealand
radians (Gmelin, 1791). New Zealand
stellifera (Gmelin, 1791). New Zealand
strigilis (Hombron & Jacquinot, 1841). Auck-
land and Campbell Ids.
subsp. bollonsi Powell, 1955. Antipodes Ids.
subsp. chathamensis (Pilsbry, 1891). Chatham
Ids.
subsp. flemingi Powell, 1955. Snares Ids.
subsp. oliveri Powell, 1955. Bounty Ids.
subsp. redimiculum (Reeve, 1854). Southern
New Zealand
+ thomsoni Powell & Bartrum, 1929. Lower Mio-
cene, New Zealand
tcophina Powell, new species. Lower Miocene,
New Zealand
taberna Powell, new species. Lower Miocene,
New Zealand 3
Genus Nacella Schumacher, 1817
Subgenus Nacella Schumacher, 1817
mytilina (Helbling, 1779). Type. Southern
Chile to Kerguelen Id.
A. W. B. Powell
kerguelenensis (E. A. Smith, 1877). Kerguelen
and Heard Ids.
Subgenus Patinigera Dall, 1905
clypeater (Lesson, 1831). Chile
concinna (Strebel, 1908). South Georgia to
Antarctica
deaurata (Gmelin, 1791). Patagonia, Falklands,
Tierra del Fuego
subsp. delicatissima (Strebel, 1907). Magellan
and Falklands
delesserti (Philippi, 1849). Marion Id.
edgari (Powell, 1957). Kerguelen Id.
flammea (Gmelin, 1791). Strait of Magellan
fuegiensis (Reeve, 1855). Magellan, Falklands,
South Georgia
magellanica (Gmelin, 1791). Type. Magellan to
Falklands
subsp. venosa (Reeve, 1854). Chiloe Island,
Chile
macquariensis Finlay, 1927.
Heard Ids.
terroris (Filhol, 1880). Campbell Id.
Macquarie and
Acknowledgements
The writer is greatly indebted to Mr. W. B.
Dixon Stroud for his continued generous mon-
etary support of this and other projects, intended
for publication, or already published, in “Indo-
Pacific Mollusca.”
To the following people who have assisted with
information, photographs, the loan of types, and
other material, the writer gratefully acknowledges
the help afforded by—Dr. R. T. Abbott, Delaware
Museum of Natural History; Professor Dr. W.
Adam, Institut Royal des Sciences Naturelles de
Belgique; Mr. W. O. Cernohorsky, Auckland In-
stitute and Museum; Dr. F. M. Climo, Dominion
Museum, Wellington; Mr. N. W. Gardner, Auck-
land; Mrs. J. Kerslake, Sydney; Dr. Y. Kondo,
B. P. Bishop Museum, Honolulu; Mr. I. G. Mar-
row, Melbourne; Dr. D. F. McMichael, formerly
Australian Museum, Sydney; Mrs. V. Orr Maes,
Academy of Natural Sciences of Philadelphia;
Dr. J. F. Peake, British Museum (Natural His-
tory); Dr. W. F. Ponder, Australian Museum,
Sydney; Mr. L. Price, Kaitaia, New Zealand; Dr.
H. A. Rehder, United States National Museum;
Dr. J. D. Taylor, British Museum (Natural His-
tory); and the late Mr. D. Thaanum, Honolulu.
To Mrs. Nancy Prior of Cape Town, the writer
is especially indebted for the fine examples of
South African limpets provided for the colour
plates.
[01-566]
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patellidae 91
Selected Bibliography
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Born, I. von. 1778. Index rerum naturalium Musei Caesarei
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Brian, M. V. and Owen, G. 1952. The Relation of the radula
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Christiaens, -J. 1967. Quelques considerations sur les Patel-
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Christiaens, J. 1968. Validite’ du nom Patella piperata Gould.
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366-373.
Christiaens, J. 1968. La Patelle bleue de Vile de Ste-Héléne.
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Cossmann, M. and Peyrot, M. A. 1917. Conchologie Neo-
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Dall, W. H. 1870. On the Limpets; with special reference to
the species of the West Coast of America, and to a more
natural classification of the group. American Journal of
Conchology, vol. 6, part 3, pp. 227-282; pls. 14-17.
Dautzenberg, Ph. and Fischer, P. H., 1925. Les Mollusques
Marins du Finistére et en particulier de la Région de Ros-
coff. Trav. Stat. Biol. Roscoff, 3, pp. 97-101 (Patellidae).
Deshayes, G. P., 1860-66. Description des Animaux sans
Vertébres découverts dans le bassin de Paris, pour servir de
supplement a la Description des Coquilles fossiles, vol. 3.
Dillwyn, L. W. 1817. A Descriptive Catalogue of Recent
Shells Arranged According to the Linnaean Method. London,
vol. 2. (Patellidae), pp. 1015-1062.
Dodd, J. M. 1956. Studies on the Biology of Limpets, 3. Her-
maphroditism in the three British species of Patella. Jour.
Mar. Biol. Assoc. U. K., vol. 35, pp. 327-340.
Eales, Nellie B., 1923. Anatomy of Gastropoda (except the
Nudibranchia), British Antarctic (“Terra Nova’) Expedition,
1910. Zoology, Mollusca, pt. 5, pp. 3-6 (Patellidae).
Evans, R. G. 1947. Studies on the Biology of British Limpets.
Proceedings of the Zoological Society, London, vol. 117,
pp. 411-423.
Fischer-Piette, E. 1935. Systematique et Biogeographie—les
Patelles d'Europe et d'Afrique du Nord. Journ. de Conchyl.,
vol. 79, pp. 5-66.
Fischer-Piette, E. 1942. Les Mollusques d’Adanson. Journ. de
Conchyl., vol. 85, no. 2, pp. 103-196; continued, no. 3, pp.
197-292; concluded, no. 4, pp. 293-377.
Fischer-Piette, E. and Gaillard, J.-M. 1959. Les Patelles au
long des cotes Atlantiques Ibetiques et Nord-Marocaines.
Journ. de Conchyl., 99 (4), pp. 135-200.
Fischer, P. H. and Kisch, B. S. 1958. Habitacles de Patelles
Creuses sur le Test d’Autres Patelles. Journ. de Conchyl.
vol. 98 (4), pp. 168-170.
Fretter, V. and Graham, A. 1962. British Prosobranch Mol-
luses. Ray Society, London, pp. 1-755. (Numerous references
to Patellidae, index, p. 753).
Gmelin, J. F. 1791. Systema naturae per regna tria naturae.
13th ed. Leipzig, 1 (6), pp. 3691-3786 (Patella).
Graham, A. 1932. On the Structure and Function of the Ali-
mentary Canal of the Limpet. Trans. Roy. Soc. Edinb. vol.
57, pp. 287-308.
Helbling, H. S. G. 1779. Beitriige zur Kenntnis neuer und
seltener Conchylien. Abh. Privatg. Bohmen, vol. 4, pp.
102-131.
Hombron, J. B. and Jacquinot, C. H., 1841. Suite de la de-
scription de quelques Mollusques, provenant de la cam-
pagne de l’Astrolabe et de la Zélée. Annales des Sciences
Naturelles, Zool. & Paleont. ser. 2, vol. 16, pp. 190-192.
Iredale, T. 1940. Marine Mollusca from Lord Howe Island,
Norfolk Island, Australia, and New Caledonia. Australian
Zoologist, vol. 9, pt. 4, pp. 432-433 (Patellidae).
Jay, J. C. 1839. Catalogue of Recent Shells in the Cabinet of
J. C. Jay, 3rd. ed., pp. 1-125, New York.
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tives of the Genus Patella Linnaeus. Annals of the Natal
Museum, vol. 11, pt. 3, pp. 487-517.
Knight, J. Brooks, et al. 1960. Moore’s Treatise on Paleontol-
ogy. Part 1, Mollusca 1, pp. 231-237 (Patellacea).
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pp. 1-140.
Lesson, R. P. 1831. Voyage Autour du Monde, éxecuté par
ordre du Roi, sur la corvette de La Majesté, La Coquille,
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Paris, vol. 2 (1), pp. 239-471.
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pp. 1-194.
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10th ed. Stockholm, pp. 1-824.
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Marine Gastropods. Science, vol. 137, No. 3526, pp. 279-
280.
Lowenstam, H. A. 1962. Magnetite in Denticle Capping in
Recent Chitons (Polyplacophora). Geol. Soc. America, Bull.
73, pp. 435-435.
Macpherson, J. Hope. 1955. Preliminary Revision of the Fam-
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Martyn, T. 1784-1786. The Universal Conchologist. London,
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(1786).
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Museum de Geneve. Mollusques vivants, 1. Revue Suisse de
Zoologie, vol. 57, no. 34, pp. 689-701 (Patella).
Michelotti, G. 1847. Description des Fossiles des Terrains
Miocenes de I'Italie septentrionale. Holl. Maat. Wetensch.
Nat. Verh., vols., 2, 3 (2). Haarlem.
Nordsieck, F. 1968. Die europaischen Meeres-Gehiusesch-
necken (Prosobranchia) Vom Eismeer Kapverden und
Mittelmeer. Stuttgart. pp. 13-15 (Patellidae).
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Orton, J. H. 1928. Observations on Patella vulgata, pt. 2:
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15, pp. 863-874.
Orton, J. H. 1929. Observations on Patella vulgata, pt. 3:
Habitats and Habits. Jour. Mar. Biol. Assoc. U. K., vol. 16,
pp. 277-288.
Pallary, P. 1912. Exploration scientifique du Maroc. Mala-
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Payraudeau, B. C. 1826. Catalogue descriptif et methodique
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[01-571]
92 Patellidae
A. W. B. Powell
Perry, G. 1811. Conchology, or the natural history of shells.
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80-84 (Patellidae).
Powell, A. W. B. 1955. Mollusca of the Southern Islands of
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(Patellidae).
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Schaffer, F. X. 1912. Das Miocan von Eggenburg. Abh. geol.
Reichs. Anst., vol. 22 (2), pp. 129-193.
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Smith, F. G. W. 1935. The Development of Patella. Phil.
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Fauna and Flora of South Africa. Part .3. Annals of the Natal
Museum, vol. 11, pt. 2, pp. 207-324.
Strebel, H. 1907. Beitrage zur kenntnis der Molluskenfauna
der Magalhaen-Provinz, No. 5, pp. 110-155; pls. 3-7 (Patel-
lidae).
Thiele, J. in Troschel & Thiele, 1891. Das Gebiss der
Schnecken, vol. 2 (7), pp. 309-334; pl. 28 (Patellidae).
Thiem, H. 1917. Beitrage zur Anatomie und Phylogenie der
Docoglossen. 1. Zur Anatomie von Helcioniscus ardosiaeus
Hombron et Jacquinot unter Bezugnahme auf die Bear-
beitung von Erich Schuster in den Zoolog. Jahrb., Supple-
ment 13, vol. 4, 1913. Jena Z. Naturw. vol. 54, pp. 333-404.
Thomson, J. A. 1919. Polymorphism in the Common New
Zealand Limpet, Cellana radians (Gmelin). N. Z. Journ. Sci.
& Tech., vol. 2, pp. 264-267.
Tomlin, J. R. le B. and Stephenson, T. A. 1942. South African
Patellidae. Proc. Malac. Soc. London, vol. 25, pt. 1, pp. 4-9.
Turton, W. H. 1932. The Marine Shells of Port Altred S. At-
rica. Oxford Univ. Press, pp. 160-171 (Patellidae).
Wood, W. 1828. Supplement to the Index Testaceologicus; or
a catalogue of shells, British and Foreign, London, pp.
1-59.
[These occasional blank areas occur between
genera and subgenera to permit the insertion
of new material and future sections in their
proper systematic sequence. |
(01-5
-~l
bo
]
November 27, 1973 INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patellidae 93
Plate 66. Patella of the subgenus Scutellastra
Fig. 1. Patella (Scutellastra) kermadecensis Pilsbry, 1894. Figs. 3-4. Patella (Scutellastra) laticostata Blainville, 1825.
Raoul Island, Kermadec Islands. Largest living species of the Albany, Western Australia.
subgenus. Fig. 5. Patella (Scutellastra) tabularis Krauss, 1848. Cape
Fig. 2. Patella (Scutellastra) flexuosa subspecies optima Point, South Africa.
Pilsbry, 1927. Yakushima, Japan.
[01-577]
94 Patella
Family Patellidae Rafinesque 1815
The Patellidae or family of true limpets have
simple, oval to rounded, conical or cap-shaped
shells, without a perforation, marginal notch or
internal septum. They are characteristic of the
intertidal zone and seldom extend much below
low-tide mark. A detailed account of the animal,
its habits, functions, distribution and geological
range, is given in the introductory section of this
work.
Subfamily Patellinae Rafinesque, 1815
The subfamily Patellinae includes the genera
Patella and Helcion, as well as several subgenera
of each.
The radula comprises four identical central
teeth, often with the addition of a median central
that may vary from vestigial to fully developed.
The lateral is large and pluricuspid, and_ is
flanked by three weak, slender, apparently func-
tionless marginals. The radular ribbon is relatively
short, straight, and folded back upon itself at the
nascent end.
The gill cordon is continuous in all members,
except in typical Helcion, which has the cordon
interrupted by the head, understandable in that
instance, since the sole species, pectunculus, has
the anterior end reduced almost to nothing.
The shell in Patella is usually rather solid, por-
cellanous within, and seldom iridescent. On the
other hand, Helcion (Patinastra) is semitrans-
parent, and Cellana-like, except for the dentition
which closely resembles that of Patella.
The typical genus, Patella, is distributed along
most of the eastern coast of Europe, from the
Lofoten Islands, and including Britain, to Spain,
the Mediterranean, west coast of Africa and off-
shore islands; also there is one species in Natal,
and another, somewhat atypical, at the South
Indian Ocean Islands of St. Paul and Amsterdam.
The subgenus Patellona is predominantly West
African, but extends to South Africa; the sub-
genera Cymbula and Olana are exclusively South
African; the subgenus Scutellastra is South A fri-
can as well, but also has a very extensive Indo-
Pacific range, and the subgenus Ancistromesus,
A. W. B. Powell
Patellidae
largest of all limpets, belongs exclusively to the
west coast of Central America.
Numerous species, attributed to Patella, Hell-
cion and Nacella, have been described from
European Cretaceous and Tertiary horizons, but
their true identity, of necessity based upon shell
characters alone, is uncertain.
Genus Patella Linnaeus, 1758
Type: Patella vulgata Linnaeus, 1758
Shell ovate, conical or cap-shaped, with the
apex subcentral, usually solid, and of medium size
to very large. Sculpture consisting of radial ridges
of varying strength, mostly crossed by concentric
growth lines. Interior of shell varying from sub-
translucent, polished and _ iridescent, to opaque
porcelanous. Colour pattern external, usually in
the form of radials associated with the ribbing,
and showing through to the interior in subtrans-
lucent shells but confined to the marginal border
in those with a thick porcelanous internal layer.
The gill cordon is complete, and the radula
relatively short and folded back upon itself at the
nascent end. The radula formula is—
341444143 oF
ot 14 (24 Le 2) 1+ sor
SPO De 3
The variations of the above formulae occur
in the central teeth, which may consist of 4 iden-
tical centrals in a horizontal row, as in Patella
vulgata, or in others when an incipient median
central is added, or, again, in certain species of
the subgenus Scutellastra when the median cen-
tral attains the size of the other centrals, thus
making 5 identical centrals. The lateral is almost
invariably pluricuspid, and the 3 marginals are
narrow, with very weak cusps at most, and they
are apparently functionless.
The range of the genus is wide-spread in warm
and temperate seas, but is absent from certain
regions, notably both coasts of North America,
the Caribbean and South America.
A number of fossil species attributed to Patella
has been described, ranging from the upper Cre-
taceous onward, but most of these are difficult to
assign generically or even to family since we lack
knowledge of the soft parts.
The genus Patella is here divided into several
subgenera that are each more or less restricted to
definite geographical areas. Their synonymy is
recorded under the relevant subgenera.
[01-578]
November 27, 1973
Subgenus Patella Linnaeus, 1758
Type: Patella vulgata Linnaeus, 1758
Shell of small to moderate size, the inner layer
subtranslucent and more or less iridescent, often
with the external colour pattern showing through
the glaze. Gill cordon complete and radula with
4 identical central teeth, arranged in a horizontal
row, and occasionally with an incipient median
central, represented by a narrow functionless
plate.
Distribution, the western coastline of Europe,
from the Lofoten Islands, and including Britain,
to the Mediterranean, down to Madeira and the
Canary Islands, and appearing again along the
coast of Natal.
Synonymy—
1758 Patella Linnaeus, Syst. Nat., ed. 10, p. 780. Type, by
subsequent designation, Fleming, 1818: Patella’ vul-
gata Linnaeus, 1755S.
1810 Patellus Montfort, Conchyliologie Systématique, vol. 2,
p. 67. Type, by original designation: Patellus roseus
Montfort, 1810.
1884 Patellopsis Thiele in’ Troschel, Das Gebiss der
Schnecken, vol. 2, p. 324, based upon the radula (pl.
28, fig. 22) of an unnamed South African Patella, pos-
sibly variabilis Krauss, 1848.
1884 Patellastra Monterosato, Natural. Sicil., vol. 3, p. 103.
Type, by monotypy: Patella lusitanica Gmelin, 1791.
1912 Costatopatella Pallary, Mem. Inst. Egypte, vol. 7 (3),
p. 148.
1920 Granopatella Pallary, Arch. Sci. Prot. Franc. Expl. Sci.
Maroc., fasc. 2, p. 72.
1920 Laevipatella Pallary, Arch. Sci. Prot. Franc. Expl. Sci.
Maroc., fase. 2, p. 72.
Patella vulgata Linnaeus, 1758
(Pl. 63, figs. 1-3; pl. 68, figs. 1, 2; pl. 61, fig. 1)
Range—Western Europe, Lofoten Islands to
Spain and the British Isles.
Remarks—This is the common European edible
limpet. It is moderately large, solid, oval and
conical, radially ribbed, and usually whitish or
yellowish, often radially lined or streaked in
brown.
Description—Shell moderately large, up to 60
mm. (2% inches) in length, solid, oval, conical,
with the apex a little in front of the middle, and
sculptured with radiating ribs and interstitial lirae.
Colour varying from whitish to yellowish,
sometimes radially lined or streaked with dark-
brown. Interior weakly iridescent, the spatula
grayish to leaden colour or clouded with whitish
INDO-PACIFIC MOLLUSCA, vol. 1, no. 15
Patella 95
callus, often with the shell margin dark-lined by
the external pattern showing through.
Radula—Formula 3+1+4+1+3. The four
central teeth are of approximately equal size, and
are arranged in a straight horizontal line, without
a median vestigial central.
Measurements (mm.)—
length width height
60.5 53.0 32.0 Caldy Island, S$. Wales
43.0 O75 21.0 Isle of Man
Synonymy—
1758 Patella vulgata Linnaeus, Syst. Nat., ed. 10, p. 752.
? 1798 Patella conus Réding, Mus. Bolten., pt. 2, p. §.
1811 Patella radiata Perry, Conch., London, pl. 43, fig. 1
(non Born, 1778).
1839 Patella conica Anton, Verzeichniss, p. 26 (non Blain-
ville, 1825).
1844 Patella vulgata var. conica Brown, Illust. Conch., ed. 2,
p. 65.
1844 Patella vulgata var. communis Brown, Illust. Conch.,
ed. 2, p. 63.
1854 Patella vulgata Linn., Reeve, Conch. Iconica, vol. 8,
pl. 18, figs. 42 a-c. (Dec.).
1857 Patella vulgata var. intermedia Knapp (in Murray), Ann.
Mag. Nat. Hist., 19, p. 211.
1865 Patella vulgata var. elevata Jeffreys, Brit. Conch., vol. 3,
p. 237.
1865 Patella vulgata var. picta Jeffreys, Brit. Conch., vol. 3,
p. 237.
1887 Patella vulgata var. secernenda Dautzenberg, Excur.
mal. St.-Lunaire, p. 13.
1891 Patella vulgata Linn., Pilsbry, Man. Conch., vol. 13,
p. 52, pl. 10, figs. 1-6.
1906 Patella vulgata var. aurea Martel in Dautzenberg &
Durouchoux, Suppl. Faun. malac. St.-Malo, p. 11.
1906 Patella vulgata var. major Dautzenberg & Durouchoux,
Suppl. Faun. malac. St.-Malo, p. 11.
Patella ferruginea Gmelin, 1791
(PI. 69, figs. 1-3)
Range—Mediterranean, from the Aegean to Spain
and North Africa.
Remarks—This species is easily recognised by its
thick shell, strong radial ribs, deeply corrugated
margin and ashen colour.
Description—Shell moderately large, up to 62
mm. (2-7/16 inches) in length, very solid, ovate,
conical, with the apex subcentral, coarsely sculp-
tured with numerous strong radial ribs, that are
rendered scabrous by concentric growth lines, and
also strongly corrugate the margin. Colour, extern-
ally dull ashen, more or less stained with pale
brown; internally, bluish white, corrugated margin
bordered in dark-brown, almost black, and the
spatula clouded with whitish callus.
Radula—Formula 3 + 1 + (2+1+2) + 1+ 3. Radula
very similar to that of caerulea, except that the
[01-579]
96‘ Patella
small slender median central is a definite tooth
bearing a small cusp.
Measurements (mm. )—
length width height
60.0 51.5 24.0 Corsica
59.0 47.0 21.0 Corsica
Synonymy—
1791 Patella ferruginea Gmelin, Syst. Nat., ed. 13, p. 3706;
based upon Martini-Chemnitz, Conch. Cab., vol. 1, pl. 8,
tig. 66.
1819 Patella luteola Lamarck, Anim. sans vert., vol. 6 (1), p. 327.
1819 Patella pyramidata Lamarck, Anim. sans vert., vol. 6 (1),
p. 327.
1826 Patella rouxii Payraudeau, Cat. Moll. Corse, p. 90.
1826 Patella lamarckii Payraudeau, Cat. Moll. Corse, p. 90.
1854 Patella costoso-plicata Reeve, Conch. Iconica, vol. 8, pl. 8,
figs. 14 a, b.
1884 Patella ferruginea var. ficarazzensis de Gregorio, Bull. Soc.
Mal. Ital., vol. 10, pp. 120-124.
1884 Patella ferruginea var. imperatoria de Gregorio, Bull. Soc.
Mal. Ital., vol. 10, pp. 120-124.
1884 Patella ferruginea var. percostata de Gregorio, Bull. Soc.
Mal. Ital., vol. 10, pp. 120-124.
1884 Patella ferruginea var. sitta de Gregorio, Bull. Soc. Mal.
Ital., vol. 10, pp. 120-124.
1891 Patella ferruginea Gmelin, Pilsbry, Man. Conch., vol. 13,
p. 81, pl. 53, figs. 1-3; pl. 17, figs. 23, 24.
1950 Patella luteola Lamarck, Mermod, Rev. Suisse Zool., vol.
57, no. 34, p. 692, fig. 3 (type).
1968 Patella ferruginea Gmelin, Nordsieck, Eur. Meeres-
Gehauseschn. Stuttgart, p. 15.
Records—CORSICA; near Bonifacio (AWBP. coll. 28388).
SPANISH MOROCCO; Melilla (AWBP. coll.); Chafarinas Is-
lands (Zafarines), 35° 10’ N., 2° 25’ E. (AWBP. coll.).
Types—The type of luteola is in the Museum
d'Histoire Naturelle de Geneve.
Patella baudonii Drouet, 1858
(Pl. 75, figs. 1, 2)
Range—Azores, Santa Maria and Pico.
Remarks—This species, which the writer has not
seen, seems to be closely allied to, if not identical
with, Patella ferruginea Gmelin, 1791. Pilsbry’s
translation of the original description follows, and
the illustrations are from Drouet’s original figures.
Description—‘Shell large, subelevated, coarsely
ribbed, plicate, solid, thick; outside greyish-green,
inside white; vertex subacute, submedian; aper-
ture oval, a little crenated.”
Measurements (mm. )—
length width height
60.0 50.0 25.0 (Drouet)
A. W. B. Powell
Patellidae
Synonymy—
1858 Patella baudonii Drouet, Moll. Mar. Agores, p. 41, pl. 2,
figs. 8, 9.
1891 Patella baudonii: Pilsbry, Man. Conch., vol. 13, p. 86, pl.
54, figs. 15, 16.
Patella rustica Linnaeus, 1758
(Pl. 69, figs. 4, 5)
Range—Atlantic coast of south west France,
Portugal, Spain, Mediterranean and Adriatic Seas.
Remarks—This species, better known by the
Gmelin name, lusitanica, is rather small, ovate-
conical, and densely sculptured with fine granular
radials. The external colour is greyish, or pale
brownish, speckled with black, and internally it is
broadly radially banded in dark puplish-brown. A
nearly related species is the narrowly-ovate piperata
from Madeira and the Cape Verde Islands.
Description—Shell rather small, up to 35 mm. (1%
inches) in length, solid, ovate, tall-conical, with the
apex slightly anterior to the middle. Sculpture con-
sisting of very numerous, closely spaced, narrow,
somewhat uneven, granulose radial riblets. Colour,
externally pale yellowish-brown to greyish, often
with the rib-granules black, internally broadly
rayed in dark-brown or blue-black on a greyish-
silvery ground. Spatula white callused, often sur-
rounded by a yellowish-brown stain.
Radula—F ormula 3 + 1 + (2+1+2) + 1+ 3. Radula
with or without a narrow median central tooth,
remaining four centrals of uniform size, and ar-
ranged in a horizontal line.
Measurements (mm.)—
length width height
35.0 28.5 17.0 Melilla, Morocco
28.0 24.0 12.55. Melilla, Morocco
Synonymy—
1758 Patella rustica Linnaeus, Syst. Nat., ed. 10, p. 783.
1791 Patella lusitanica Gmelin, Syst. Nat., ed. 13, p. 3715.
1798 Patella squamata Roding, Mus. Bolten, pt. 2, p. 10.
1819 Patella punctata Lamarck, Anim. sans vert., vol. 6, p. 333.
1825 Patella subgranularis Blainville, Dict. Sci. Nat., vol. 38,
p. 113. (fide Christiaens, 1968, p. 367).
1854 Patella nigro-punctata Reeve, Conch. Iconica, vol. 8, pl.
23, figs. 57 a-c (Dec.).
1883 Patella lusitanica var. minor Marion, Faune bass. med.,
p. 48.
1884 Patellastra lusitanica Gmel., Monterosato, Natural. Sicil.,
vol. 3, p. 103.
1891 Patellastra lusitanica Gmel., Thiele, in Troschel & Thiele,
Das Gebiss der Schnecken, 2, pl. 28, fig. 12 (radula).
1891 Patella lusitanica Gmel., Pilsbry, Man. Conch., vol. 13,
p. 87, pl. 11, figs. 15-19.
1912 Patella rustica Linné var. major Pallary, Explor. scient.
Maroc., p. 72.
[01-580]
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 97
Plate 67. Indo-Pacitic Cellana
Figs. 1, 2. Cellana testudinaria (Linnaeus, 1758). Vanualava,
Banks Islands.
Fig. 3. Cellana talcosa (Gould, 1846). Molokai, Hawaiian
Islands.
Figs. 4-6. Cellana exarata (Reeve, 1854). Molokai, Hawaiian
Islands.
Fig. 7. Cellana grata (Gould, 1859). Matsushima Island,
Korea.
Figs. 8,9. Cellana radiata (Born, 1778). Colombo, Ceylon.
Figs. 10, 11. Cellana radiata subspecies capensis (Gmelin,
1791). Near Durban, Natal.
Figs. 12, 13. Cellana radiata subspecies orientalis (Pilsbry,
1891). Fig. 12. Russell Islands, Solomon Islands (note the
strong radial folds). Fig. 13. Tau Island, Samoa.
Fig. 14. Cellana radiata subspecies enneagona (Reeve, 1854).
Jolo, Philippine Islands.
Figs. 15, 16. Cellana nigrolineata (Reeve, 1854). Fig. 15.
Fukura, Awaji, Japan. Fig. 16. Chiringashima, Japan.
Figs. 17, 18. Cellana mazatlandica (Sowerby, 1539). Bonin
Islands.
[01-581]
98 Subgenus Patella
A. W. B. Powell
Patellidae
Plate 68. Figs. 1, 2. Patella vulgata Linnaeus, 1758. Fig. 1.
Kimmeridge, England, 52 mm., AWBP coll. 11358. Fig. 2.
Isle of Man, 38 mm., AWBP coll. 11359. Figs. 3, 4. Patella
aspera Roding, 1798, Caldy Island, Wales, 47 mm., AWBP
coll. 217.
1912 Patella rustica Linneé var. maroccana Pallary, Explor. scient.
Maroc., p. 72.
1938 Patella lusitanica var. orientalis Pallary, Jour. Conchyl.,
vol. 82, p. 47.
1950 Patella punctata Lam., Mermod, Rev. Suisse Zool., vol.
57 (34), p. 695, fig. 7 (type).
1968 Patella lusitanica Gmelin, Christiaens, Bull. Mus. Nat.
WHist. Nat., ser. 2, vol. 40 (2), pp. 366, 367.
1968 Patella (Patellastra) rustica L., Nordsieck, Die europ-
Meeres Gehauseschnecken, Stuttgart, p. 15.
Patella piperata Gould, 1846
(Pl. 71)
Range—Madeira and Cape Verde Islands.
Remarks—Shell very similar to that of rustica in
sculpture and in coloration, but more elongate-ovate
in its younger stages, and with the apex nearer to
the anterior end.
Radula—The radula differs from that of rustica in
that the four central teeth are not in horizontal
alignment, the middle pair being set lower than the
outer pair (Christiaens, 1968, p. 370, fig. 2a).
Plate 69. Figs. 1-3. Patella ferruginea Gmelin, 1791. Fig. 1.
Chafarines Islands, Morocco, 55 mm., AWBP. coll. 1054.
Fig. 2. Bonifacio, Corsica, 60 mm., AWBP coll. 28388. Fig. 3.
Melilla, Morocco, 42 mm., AWBP coll. 30974. Figs. 4, 5.
Patella rustica Linnaeus, 1758, Oran, Algeria, 25-26 mm.,
AWBP coll. 80.
Measurements (mm.)—
length width height
44.0 40.0 — Christiaens, 1968, p. 372
27.0 21.0 12.0 Madeira
Synonymy—
1839 Patella guttata Orbigny, in Webb & Berthelot, Hist. Nat.
Moll. Canaries, p. 98 (non Gmelin, 1791).
1846 Patella piperata Gould, Proc. Boston Soc. Nat. Hist., vol. 2,
p. 150.
1846 Patella nigrosquamosa Dunker, Zeitschr. f. Malak., p. 25.
1866 Patella frauenfeldi Dunker, Verh. k. k. zool.-bot. Ges. Wien,
vol. 16, p. 914. “Madras” in error for Madeira.
1867 Patella frauenfeldi Dunker, Frauenfeld, Reise Novara,
Zool., vol. 2, pt. 3, Moll., p. 15, pl. 2, figs. 26 a, b.
1968 Patella piperata watsoni Christiaens, Bull. Mus. Nat.
@Hist. Nat. ser. 2, vol. 40, no. 2, p. 371, text fig. 2 b;
pl. 1, fig. b.
1968 Patella piperata nigro-radiata Christiaens, Bull. Mus. Nat.
@Hist. Nat. ser. 2, vol. 40, no. 2, p. 371, text fig. 2 c; pl.
1, fig. c.
1968 Patella piperata alba Christiaens, Bull. Mus. Nat. d’Hist.
Nat. ser. 2, vol. 40, no. 2, p. 371, pl. 1, fig. g (non P. alba
Anton, 1539).
[01-582]
November 27, 1973
Patella caerulea Linnaeus, 1758
(Pls. 61, 63, 72, 74)
Range—Mediterranean and Adriatic Seas, Por-
tugal, Azores, Madeira and Canary Islands.
Remarks—This species is variable in shape, col-
our, and strength of the radial ribbing, but in gen-
eral terms it is a depressed, thin, and spreading
shell, with 6 or 7 distinct marginal angles, result-
ant from 7 to 9 prominent radial folds. The typical
form of the species from the Mediterranean and
Adriatic Seas, has a colour range, varying from al-
most white to buff or pale brownish, often radially
banded with iridescent blue.
Shells from the Azores and Madeira are larger,
even more depressed than the typical species, have
broader and more prominent radial folds, and are
of much darker colour, being dark reddish brown
externally, similarly coloured internally, but dif-
fused with iridescent blue, and with a distinct-
edged, white spatula. This latter form is crenata
Gmelin, and when more material is studied,
Gmelin’s name may prove to be usable to define a
regional subspecies of caerulea, restricted to the
Azores, Madeira and Canary Islands. Negating
this possibility is the fact that shells from the ad-
jacent mainland of Spanish Morocco have the dark
colouring of crenata but a shape and sculpture
similar to those in typical caerulea.
Description—Shell of moderately large size, 40-71
mm. (1/2-2% inches) in length, thin, depressed, usu-
ally distinctly 6 or 7 angled, resultant from 7 to 9
broadly rounded primary folds that project at the
margin. Surface crowded with secondary radials of
varying sizes, mostly imbricated by concentric
growth lines. Colour whitish or buff externally, the
interior silvery-white, radially lined or banded in
blue, the spatula bluish or white-callused. Some
examples have a pale yellowish interior without
radial markings, and the form crenata is dark red-
dish brown, internally diffused with iridescent blue,
and with a clear-cut white spatula.
Radula—Formula 3 + 1 + (2+X+2) + 1+ 3. The
radula is of the same style as that of vulgata, except
for a slight median gap between the pairs of four
centrals, in which appears a narrow vestigial plate.
The four functional centrals, as in vulgata, are in a
straight horizontal row.
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patella 99
Measurements (mm.)—(all A. W. B. Powell
collection.)
length width height
51.0 46.0 12.0 Capri, Italy
36.0 30.0 10.5 Melilla, Morocco
71.0 64.0 14.0 Madeira (crenata form)
Synonymy—
1758 Patella caerulea Linnaeus, Syst. Nat., ed. 10, p. 782.
1791 Patella crenata Gmelin, Syst. Nat., ed. 13, p. 3706.
1791 Patella margaritacea Gmelin, Syst. Nat., ed. 13, p. 3707.
1793 Patella tarentina von Salis, Reise ins. Koenig. Neapel, p.
359, pl. 6, fig. 2.
1798 Patella silicina Riding, Mus. Bolten., pt. 2, p. 9.
1819 Patella tarentina Lamarck, Anim. sans vert., vol. 6, p. 332.
1826 Patella bonnardii Payraudeau, Moll. de Corse, p. 89.
1836 Patella fragilis Philippi, Enum. Moll. Sicil., vol. 1, p. 110.
1838 Patella subplana Potiez & Michaud. Gal. Moll. Douai, vol.
1, p. 524.
1854 Patella scutellaris Lam., Reeve, Conch. Iconica, vol. 8,
pl. 20, fig. 49.
1882 Patella stellata Bucquoy, Dautzenberg & Dollfus, Moll. mar.
Roussillon. (non Helbling, 1779).
1882 Patella adspersa Bucquoy, Dautzenberg & Dollfus. Moll.
mar. Roussillon.
1882 Patella caerulea var. cognata Bucquoy, Dautzenberg &
Dollfus, Moll. mar. Roussilon, p. 471.
1882 Patella caerulea var. intermedia Bucquoy, Dautzenberg &
Dollfus, Moll. mar. Roussilon, p. 471.
1891 Patella caerulea Linne~ Pilsbry, Man. Conch., vol. 13,
p. 83, pl. 10, figs. 7-12.
1950. Patella tarentina Lam., Mermod, Rev. Suisse Zool., vol. 57
(34), p. 695 (text figs. of type series).
Records—ITALY: Naples; Isle of Capri; Palermo, Sicily. MAL-
TA. MOROCCO: Melilla. MADEIRA (crenata form). (All AWBP
coll.)
Patella moreleti Drouet, 1858
(Pl. 75, figs. 3, 4)
Range—Fayal, Azores.
Remarks—The writer has not seen this species
which possibly may be only a form of Patella
caerulea Linnaeus. Pilsbry’s translation of the orig-
inal description follows, accompanied by copies of
Drouet’s figures.
Description—‘Shell subdepressed, very rugose,
ribbed, the ribs scaly, scarcely solid; brownish-
green outside; inside brownish or reddish, irides-
cent, with a white spot at the summit. Apex acute.
Aperture ovate, crenulated.”
Measurements (mm.)—
length width height
40.0 30.0 12.0 (Drouet)
Synonymy—
1858 Patella moreleti Drouet, Moll. Mar. Agores, p. 42, pli 2,
figs. 10, 11.
1891 Patella moreleti Drouet, Pilsbry, Man. Conch., vol. 13, p.
85, pl. 56, figs. 27, 25.
[01-583]
100 Subgenus Patella
Plate 71. Patella piperata Gould, 1846. Madeira, 27 mm.,
AWBP coll. 1058.
Patella lowei Orbigny, 1839
(Pl. 74, figs. 1, 2)
Range—Canary Islands.
Remarks—This species appears to be closely
allied to the Mediterranean caerulea, from which it
differs mainly in having stronger, and more regu-
lar sculpture, resulting in a more even denticula-
tion of the margin, as opposed to the 6 or 7 distinct
marginal angles of caerulea. Also, the coloration in
lowei is darker, the exterior being rusty-brown,
and the interior dark bluish to reddish brown at the
edges, reflecting iridescent blue, and always with
a clearcut white spatula.
Description—Shell of moderate size, up to 56.5
mm. (24 inches) in length, ovate, depressed, with
the apex towards the anterior third, solid but not
thick, densely sculptured with broadly
rounded primary radials and narrow intermediates.
The margin is strongly and regularly corrugated,
the projections compound and foliated. Colour as
described above.
Measurements (mm.)—(both A. W. B. Powell col-
lection).
very
length width height
56.5 45.0 13.0 ‘Teneriffe
54.5 44.5 11.0 Teneriffe
Synonymy—
1839 Patella lowei Orbigny, in Webb and Berthelot, Hist. Nat.
Canaries, Moll., vol. 2, p. 97, pl. 7, figs. 9, 10.
1839 Patella azorica Nuttall, in Jay, Cat. Shells, ed. 3, p. 38.
1S91 Patella caerulea var. lowei Orbigny, Pilsbry, Man. Conch.,
vol. 13, p. 84, pl. 29, figs. 44-46; pl. 53, figs. 7-11.
Records—CANARY ISLANDS (Orbigny); Teneriffe (AWBP
coll. 5268).
A. W. B. Powell
Patellidae
Plate 72. Figs. 1, 2. Patella caerulea Linnaeus, 1758, Isle of
Capri, Italy, 51 mm., AWBP coll. 211. Figs. 3, 4. Patella
safiana Lamarck, 1819, Oran, Algeria, 65 mm., AWBP coll.
1959.
Patella gomesii Drouet, 1858
(PI. 74, figs. 5, 6)
Range—Azores, Bay of San Lourenzo, Santa
Maria and Pico.
Remarks—The writer has not seen examples of
this shell, which may prove to be a form of lowei.
Its distinctive character is in having about 14 very
prominent rounded radial folds, the whole surface,
folds included, being densely radially lirate.
Description—(Pilsbry’s translation of original):
“Shell large, subdepressed, rugose, ribbed-plicate,
rather solid; outside grayish-brown or rufescent;
inside shining, brown, pearly; apex situated at the
front third of the length, obtuse; aperture oval,
entire.”
Measurements (mm.)—
length width height
50-60 50-53 12-15
Synonymy—
1858 Patella gomesii Drouet, Moll. Mar. Iles Agores, p. 39,
pl. 1, figs. 6, 7.
1891 Patella gomesii Drouet, Pilsbry, Man. Conch., vol. 13, p.
86, pl. 54, figs. 17, 18.
[01-584]
November 27, 1973
Plate 73. Antarctic Australian Cellana and
INDO PACIFIC MOLLUSCA, vol. 1, no. 15 Patella 101
(for plate 70, see p. 105)
Subantarctic Nacella
Figs. 1-3. Cellana framoserica (Holten, 1802). Fig. 1. South
Australia. Fig. 2. Caloundra, Queensland. Fig. 3. Torquay,
Victoria.
Figs. 4-6. Cellana solida (Blainville, 1825). Figs. 4, 5. Stan-
ey, Tasmania. Fig. 6. South Australia (rubraurantiaca
orm).
Figs. 7, 8. Cellana ardosiaea (Hombron & Jacquinot, 1841).
Island of Juan Fernandez.
Fig. 9. Nacella mytilina (Helbling, 1779). Falkland Islands.
Fig. 10. Nacella kerguelenensis (E. A. Smith, 1877). Heard
Island. 7 -
Fig. 11. Nacella (Patinigera) deaurata (Gmelin, 1791). Falk-
land Islands.
Fig. 12. Nacella (Patinigera) terroris (Filhol, 1880). Campbell
Island. a
Fig. 13. Nacella (Patinigera) clypeater (Lesson, 1831). Chile.
Figs. 14, 15. Nacella (Patinigera) magellanica (Gmelin, 1791).
Possession Bay, Patagonia.
[01-585]
102. Subgenus Patella A. W. B. Powell Patellidae
[These occasional blank areas occur between
genera and subgenera to permit the insertion
of new material and future sections in their
proper systematic sequence. |
[01-586]
November 27, 1973
Subgenus Uncertain
The following six species of Patella are insuffi-
ciently understood, particularly with regards to their
soft anatomy, to be assigned as yet to their proper
subgenera.
Patella candei Orbigny, 1839
(PI. 75, figs. 7, 8)
Range—Canary Islands.
Remarks—The writer has not seen this species,
but from published information it appears to be
closely allied to citrullus from Funchal. Madeira.
The surface has subobsolete radials crossed by
prominent wavy concentric lirations that hap-
hazardly anastomose to form an irregular netted
appearance.
Description—(Pilsbry’s 1891 translation of the
original description): “Shell elevated, conical, thick,
smooth or irregularly roughened; ovate, margin en-
tire. Inside buff, bluish in the middle, outside pale
yellow.”
Measurements (mm.)—
length width height
67.0 55.0 27.0
Synonymy—
1839 Patella candei Orbigny, in Webb and Berthelot, Hist. Nat.
Canaries, vol. 2, Moll., p. 98, pl. 7, figs. 11, 12.
1854 Patella candei Orbigny, Reeve, Conch. Iconica, vol. 8,
pl. 15, figs. 34 a, b.
1891 Patella candei Orbigny, Pilsbry, Man. Conch., vol. 13, p.
86, pl. 55, figs. 22-24.
Patella citrullus Gould, 1846
(Pl. 75, figs. 9, 10)
Range—Funchal, Madeira.
Remarks—The writer has not seen examples of
this species which appears to be related to candei.
Pilsbry (1891, l.c.) remarked that the external sur-
face resembles the skin of a cucumber.
Description— (original) “Shell sub-diaphanous,
thin sub-conical, moderately elevated, summit
prominent; apex anterior, acute, feebly incurved,
usually somewhat eroded; a great number of faintly
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patella = 103
elevated lines, studded with fine tubercles or asper-
ities, radiate from it, and become obsolete about
half way towards the margin. Striae of increment
coarse and irregular, overlaying each other, so as
to give the shell a rude, concentrically squamose
aspect externally; disc nearly oval, a little narrowed
anteriorly; margin very thin and sharp, finely and
irregularly undulated. External colour a dusky
olive-green, with a shade of brown showing through
it, ornamented with concentric, undulating lines of
obscure white. Interior greenish-white, with bright
iridescent reflections; a slight spatulaform deposit
at the fundus, bluish at the edges and forepart,
passing into greenish towards the middle and pos-
terior portions.”
Measurements (mm.)—
length width height
45.0 32.0 7.0
Synonymy—
1846 Patella citrullus Gould, Proc. Boston Soc. Nat. Hist., vol.
2, p. 149.
1891 Patella citrullus Gould, Pilsbry, Man. Conch., vol. 13, p. 86,
pl. 28, figs. 39-41.
1964 Patella citrullus Gould, Johnson, U.
239, p. 56.
S. Nat. Mus. Bull.
Patella concolor Krauss, 1848
(Pls. 64, 76, 78)
Range—Natal coast to as far south and west as
Bushman’s River, near Port Elizabeth.
Remarks—The former name of this well-known
South African limpet, Patella variabilis Krauss,
1848, is invalidated by two prior homonyms, those
of Roding, 1798, and Risso, 1826. The earliest valid
name to replace variabilis is concolor which is the
uniformly dark-ashen colour form of this species.
The species is exceedingly variable in colour pat-
tern, but the shape, which is ovate, distinctly nar-
rowed in front, remains constant. Also it is of light
build and is often semi-transparent.
Description—Shell rather small, usually between
30 and 35 mm. in length, but occasionally attaining
50 mm. (2 inches) in length, of light build, some-
times semi-transparent, rather depressed, ovate, but
distinctly narrowed at the anterior end. Sculptured
with about 80 fine but somewhat unequal radial
ribs, crossed by dense inconspicuous concentric
lirae. Apex subcentral to about the anterior third,
the area in its vicinity usually smooth. Colour ex-
extremely variable, ranging from plain yellow, pale
yellowish brown, and rusty-brown (concolor) to al-
most black, and variously maculated; sometimes the
[01-597]
104. Subgenus uncertain
A. W. B. Powell
Patellidae
yellow form has one, or several, dark-brown radial
streaks, and the black form (polygramma) has the
primary radials picked out in white; the common
form is pale yellowish brown, radially lined and
speckled in dark-brown; spatula ill-defined, light
brownish or clouded with white callus.
Radula—Formula 3 + 1 + (2+0+2) + 1 + 3. The
radula differs from those of all other South African
patellids in the absence of the middle member of
the central teeth; the four remaining centrals, how-
ever, are grouped in pairs with a space between
them, whereas in the radula of Patella vulgata and
other European patellids, the four centrals are
closely grouped, without space for a middle mem-
ber.
Plate 74. Figs. 1, 2. Patella lowei dOrbigny, 1839, Teneriffe,
Canary Islands, 56.5 mm., AWBP coll. 5268. Figs. 3, 4. Pa-
tella ct. caerulea Linnaeus, 1758 (crenata form), Madeira,
70 mm., AWBP coll. 675. Figs. 5, 6. Patella gomesii Drouet,
1858, Azores, 50-60 mm. From Pilsbry, 1591, pl. 54, figs.
17, 18.
Measurements (mm.)—
length width height
50.0 45.0 11.25 South Africa
48.5 41.25 14.00 Natal coast
35.0 30.5 9.00 Port Alfred
28.5 24.0 7.00 Coffee Bay
Synonymy—
1848 Patella variabilis Krauss, Sudafr. Moll., Stuttgart, p. 55, pl.
3, fig. 12 (non P. variabilis Risso, 1826).
1848 Patella variabilis var. fasciata Krauss, Sudafr. Moll., Stutt-
gart, p. 55, pl. 3, fig. 12 a (non P. fasciata Gmelin, 1791).
1848 Patella variabilis var. radiata Krauss, Sudafr. Moll., Stutt-
gart, p. 59, pl. 3, fig. 12 b. (non P. radiata Born, 1778).
1548 Patella variabilis var. concolor Krauss, Sudafr. Moll.,
Stuttgart, p. 55, pl. 3, fig. 12 c.
1891 Helcioniscus variabilis Krauss, Pilsbry, Man. Conch., vol.
13, p. 147, pl. 16, figs. 18-20.
Plate 75. Figs. 1, 2. Patella baudonii Drouet, 1858, Azores,
60 mm. Figs. 3, 4. Patella moreleti Drouet, 1858, Fayal,
Azores, 40 mm. Figs. 5, 6. Patella rangiana Rochebrune,
1882, Cape Verde Islands, 44 mm. Figs. 7, 8. Patella candei
d@Orbigny, 1839, Canary Islands, 67 mm. Figs. 9, 10. Patella
citrullus Gould, 1846, Funchal, Madeira, 45 mm. (All figures
from Pilsbry, 1891, Manual of Conchology, vol. 13, plates
45, 54, 55, 56 and 58).
[01-598]
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 1, no. 15 Patella
105
Plate 70. New Zealand Cellana
Figs. 1-4. Cellana radians (Gmelin, 1791). Fig. 1. Mount
Maunganui, Bay of Plenty (earlii pattern). Figs. 2, 3. Mo-
tuihi Island, Auckland. Fig. 4. Herekopare Island, Stewart
Island (perana form).
Figs. 5, 6. Cellana flava (Hutton, 1873). Fig. 5. East Cape.
Fig. 6. Cape Campbell, Marlborough.
Figs. 7, 8. Cellana denticulata (Martyn, 1784). Mount Maun-
ganui, Bay of Plenty.
Figs. 9-11. Cellana stellifera (Gmelin, 1791). Fig. 9. Whan-
(for plate 73, see p. 101)
garei Heads. Fig. 10. Ti Point, Hauraki Gulf. Fig. 11. Long
Beach, Bay of Islands (bleached coloration of beach shells).
Figs. 12, 13. Cellana ornata (Dillwyn, 1817). Fig. 12. Mo-
tutara, West Coast, Auckland. Fig. 13. Mount Maunganui.
Figs. 14-16. Cellana strigilis (Hombron & Jacquinot, 1841).
Campbell Island.
Figs. 17-19. Cellana strigilis subspecies redimiculum (Reeve,
1854). Kartigi Beach, North Otago, South Island, New
Zealand.
[01-599]
106 Subgenus uncertain
A. W. B. Powell
Patellidae
1921 Patella variabilis constellata G. B. Sowerby, Proc. Malac.
Soc., Lond., vol. 14, p. 127.
1931 Patella variabilis Krauss, Tomlin, Ann. Natal Mus., vol. 6
(3), p. 417.
1931 Patella variabilis fasciolata Tomlin, Ann. Natal Mus., vol.
6 (3), p. 418; nom. nov. pro P. variabilis fasciata Krauss,
1848 (non Gmelin, 1791).
1931 Patella variabilis polygramma Tomlin, Ann. Natal Mus.,
vol. 6 (3), p. 418; nom. nov. pro P. variabilis radiata
Krauss, 1848 (non Born. 1778).
1932 Patella variabilis Krauss, Turton, Mar. Shells Port Alfred,
p. 167, sp. 1187.
1932 Patella variabilis fasciata Krauss, Turton, Mar. Shells.
Port Alfred, p. 167, sp. 1188.
1932 Patella variabilis radiata Krauss, Turton, Mar. Shells Port
Alfred, p. 167, sp. 1189.
1932 Patella variabilis concolor Krauss, Turton, Mar. Shells Port
Alfred, p. 168, sp. 1190.
1932 Patella variabilis constellata Sby., Turton, Mar. Shells Port
Alfred, p. 168, sp. 1191.
1932 Patella variabilis helvola Turton, Mar. Shells Port Alfred,
p. 168, sp. 1192.
1932 Patella rietensis Turton, Mar. Shells Port Alfred, p. 167,
pl. 38, fig. 1183.
1932 Patella rota (non Gmelin, 1791) Turton, Mar. Shells Port
Alfred, p. 168, sp. 1193.
1932 Patella helena Turton, Mar. Shells Port Alfred, p. 168, pl.
39, fig. 1194.
1932 Patella conspicua (non Philippi, 1849) Turton, Mar. Shells
Port Alfred, p. 168, sp. 1196 (in part).
1932 Patella farquhari Turton, Mar. Shells Port Alfred, p. 170,
pl. 40, fig. 1207.
1949 Patella variabilis Krauss, Koch, Ann. Natal Mus., vol. 11
(3), p. 510, pl. 23, figs. 1-11; text figs. 21, 22 (radula).
Records—SOUTH AFRICA: Natal coast to as far south and
west as Port Elizabeth (Koch, 1949); Natal (ex Koch; AWBP
coll.); Umtwalumi, 22 miles N. of Port Shepstone (V. Orr, 1955;
ANSP); Port St. John’s Pondoland (V. Orr; ANSP); Coffee
Bay, Transkei (V. Orr, 1955; ANSP); Port Alfred (USNM);
(AWBP coll.); near Durban (Mrs. N. Prior).
Patella depsta Reeve, 1855
(PI. 77; pl. 78, fig. 1)
Range—Islands of St. Paul and Amsterdam, South
Indian Ocean.
Remarks—Reeve cited “Macao and the Island of
St. Paul” as localities for this species, but the first
mentioned location is obviously a mistake. St. Paul
is here nominated as the type locality. The species
also occurs at the adjacent island of Amsterdam.
Gaillard (1954) figured the radula of depsta, and
assigned the species to Cellana, but the radula sug-
gests much closer alliance with the Patellinae, and
except for the laterals, is not unlike that of Patella
(Patellona). The laterals in. the Patellinae are usu-
ally fused at the base, and have a pluricuspid head,
but Gaillard’s drawing shows a pair of laterals on
either side, each separated at the base. Since the
writer has no preserved material of this species the
Plate 76. Figs. 1-7. Patella concolor Krauss, 1548. Figs. 1, 2.
Port Alfred, South Africa, 35-36 mm., AWBP. coll. 30872;
227788. Figs. 3, 4. Port St. Johns, Pondoland, South Africa,
26-34 mm., AWBP coll. 30822. Fig. 5. Coffee Bay, Transkei,
South Africa, 30 mm., AWBP coll. 48225. Fig. 6. Umtwalumi,
South Africa, 34 mm., AWBP coll. 211735. Fig. 7. South
Africa, 50 mm., AWBP coll. 46130.
[01-600]
November 27, 1973
Plate 77. Patella depsta Reeve, 1854. Island of St. Paul, South
Indian Ocean, 32-35 mm., AWBP coll. 46133.
apparently unusual form of the laterals cannot be
confirmed at present.
Description—Shell of moderate size, up to 35.5
mm. (1% inches) in length, lightly built, ovate,
gradually narrowed in front, and moderately ele-
vated, with the apex anterior to the middle, com-
pressed and hooked. Sculptured finely and deli-
cately radially lirate, arranged more or less in
fours, the inner two weaker than the outer two,
and about 100 lirae in all. Colour of exterior pale
pinkish chestnut; interior orange-brown with a
slight bronzy sheen; spatula pinkish white.
Plate 78. Fig. 1. Patella depsta Reeve, Island of St. Paul.
Radula, from Gaillard, 1954, p. 521, fig. 1. Fig. 2. Patella
concolor Krauss, Natal. Radula, from Koch, 1949, p. 511,
fig. 22 (as variabilis Krauss).
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patella 107
Measurements (mm.)—(both A. W. B. Powell
collection).
length width height
35.5 28.0 14.0 St. Paul
32.25 24.0 12.0 St. Paul
Radula—Formula 3 + 2? + (2+1+2) + 2? + 3.
Synonymy—
1855 Patella depsta Reeve, Conch. Iconica, pl. 31, figs. 85 a, b.
(Jan.).
1891 Helcioniscus depsta Reeve, Pilsbry, Man. Conch., vol. 13,
p. 151, pl. 20, figs. 45, 46.
1954 Cellana devsta Reeve, Gaillard, Bull. Mus. Nat. d’Hist.
Nat., vol. 26, pp. 520, 521, text. fig. 1 (radula).
Patella rangiana Rochebrune, 1882
(Pl. 75, figs. 5, 6)
Range—Porto Praya, Cape Verde Islands.
Remarks—This species, which the writer has not
seen, appears to be a distinctive one, with its very
scaly prominent radial ribs. Pilsbry’s translation of
the original description follows, accompanied by
copies of Rochebrune’s figures.
Description—‘“Shell ovate, depressed-convex,
rufous; vertex submucronate, usually eroded,
situated at 2/3 of the length; having larger and
smaller radiating broad, very scaly ribs, scales
subimbricating, obtuse, lenticular; margin un-
dulating; interior bluish, silvery-pearly, rayed with
bands and spots of purplish, the center spatulate.”
Measurements (mm.)—
length width height
44.0 36.0 19.0 (Rochebrune)
Synonymy—
1882 Patella rangiana Rochebrune, Bull. Soc. Philomathique,
Paris, ser. 7, vol. 6, p. 29.
1891 Patella rangiana Rochebrune, Pilsbry, Man. of Conch.,
vol. 13, p. 89, pl. 58, figs. 42, 43.
? Patella kaffraria Rennie, 1930
(PI. 79)
Range— Upper Cretaceous of Pondoland, South
Africa.
Remarks—The author of this species remarked
that “It need hardly be stated that the genus
Patella is here used in the widest possible sense.
The species is apparently distinct from any pre-
viously described from the Cretaceous.” It certainly
appears to belong to the Patellacea, but a precise
generic or even familial allocation would be purely
conjectural. Rennie’s original description follows.
[01-601]
108 Subgenus uncertain
Plate 79. PPatella kaffraria Rennie, 1930. Upper Cretaceous
of Pondoland, South Africa, 32.56 mm. Holotype, from Rennie,
1930, pl. 24, figs. 1, 2.
Description—“Shell moderately convex, with
the apex obtusely pointed, not recurved, and
placed well in front of the middle; the sides are
straight, or only slightly convex. Aperture oval,
A. W. B. Powell
Patellidae
considerably longer than wide, with wavy margin.
Surface with stout, rather irregular, radial ribs,
and narrow furrows; on the posterior side the ribs
are of two sizes, the larger and smaller alternat-
ing; on the anterior side there are more numer-
ous, finer ribs; the ribs are crossed by irregular
growth markings.”
Measurements (mm.) (Not stated, but evidently
the figures are natural size)—
length width height
32.5 25.0 14.0 holotype
Synonymy—
1930 Patella kaffraria Rennie, Annals of South African Mu-
seum, vol. 28, p. 206, pl. 24, figs. 1-4.
Types— The holotype (No. 8477) and paratype (No.
8572) are in the South African Museum.
(01-602]
November 27, 1973
Subgenus Patellona Thiele in Troschel, 1891
Type: Patella granatina Linnaeus, 1758
This group of patellids was named because of a
marked difference in the radula from that of typ-
ical Patella. The four central teeth of true Patella
occur in a horizontal alignment whereas in Patel-
lona there is a median central, narrower and of
smaller size than the outer pairs of centrals, which
instead of being in line, descend steeply to the
laterals, their tops forming a chevron. The cusps
of the centrals and laterals vary between oblique
heart-shape and parrot-beaklike.
This chevron-like radula is found in species
from Cape Verde Islands, Senegal, Guinea, An-
gola and St. Helena, as well as South Africa,
where the type species granatina and the related
oculus occur, these two being more or less re-
stricted to the cooler waters of the west coast.
Shells of this subgenus are slightly iridescent
within and the shell substance is sufficiently
transparent for the external colour patterns to
show through faintly.
The northward flowing cool Benguela Current
could account for the presence of the subgenus in
Angola and St. Helena, but locations north of
there, in the tropical waters of West Africa and
the Cape Verde Islands, are, under present con-
ditions, out of range of the influence of that cur-
rent. Nevertheless the style of radula in the trop-
ical West African and Cape Verde Islands limpets
is so similar to that of the cool water species of
the South African west coast that some distribu-
tional continuity, under more uniform hydro-
logical conditions, must have existed in the past.
Related to Patellona is the subgenus Cymbula
(see ahead) in which the central teeth have the
same chevronlike alignment, but their cusps are
distinctive in having broad blunt tops with raised
marginal rims.
Synonymy—
1891 Patellona Thiele in Troschel, Das Gebiss der Schnecken,
vol. 2, p. 317, for granatina Linnaeus, 1758, adansonii
Dunker, 1853 and plumbea Lamarck, 1819. Type, by
subsequent designation, Tomlin, 1931: Patella grana-
tina Linnaeus, 1758S.
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patella = 109
Patella granatina Linnaeus, 1758
(Pl. 64, figs. 1-3; pl. 80; pl. 82, fig. 2)
Range—South Africa, Port Nolloth on the west
coast, south to False Bay and extending east-
ward to Danger Point.
Remarks—This large but relatively thin South
African limpet is easily recognised by its broadly
ovate, almost pentagonal outline, strong, nar-
rowly crested, radial ribs, and distinctive colora-
tion of the interior which is bluish white with a
clearly outlined dark-brown spatula.
Description—Shell large, up to 85 mm. (3%
inches) in length, strong but of relatively light
build, broadly ovate and _ tall-conical, with the
apex almost central. Sculpture of radiate folds
that strongly corrugate the margin; five of the
radials on the posterior half of the shell are
stronger than the rest; radials and interspace
alike bear closely-spaced cords that are rendered
scabrous by dense concentric growth-lamellae.
Colour of exterior greyish to dull-white with an
underlying pattern of dark-brown, zigzag, con-
centric markings, often forming a netted design;
interior bluish white, the spatula dark-chocolate,
with clearly defined edges, and a marginal pat-
tern of numerous short, dark-brown dashes, with
regular gaps corresponding to the external pri-
mary radials.
Radula—Formula 3+ 1 + (2+1+2) + 1+ 3. The
median central is small and slender, flanked by a
pair of stout fully-developed centrals on either
side, followed by a pluricuspid lateral, and the
usual three, more or less functionless, marginals.
The centrals, collectively, form a chevron, as in
other members of this subgenus. The cusps of the
paired centrals and the pluricuspid laterals are
leaf-shaped, obliquely flexed, and with a median
groove or depression.
Plate 80. Patella (Patellona) granatina Linnaeus, 1758. Table
Bay, South Africa, 71 mm., AWBP coll. 183.
[01-613]
110 Subgenus Patellona
A. W. B. Powell
Patellidae
Measurements (mm.)—(all A. W. B. Powell
collection)
length width height
85.0 78.5 30.0 South Africa
74.0 63.5 35.0 South Africa
60.0 49.5 20.0 False Bay
Synonymy—
1758 Patella granatina Linnaeus, Syst. Nat. ed. 10, p. 782.
1819 Patella apicina Lamarck, Anim. sans vert., vol. 6 (1),
p. 324.
1848 Patella granatina Lam., Krauss, Sudafr. Moll., Stuttgart,
p. 43.
1854 Patella granatina Lam., Reeve, Conch. Iconica, pl. 3,
figs. 4a, b.
1891 Patella (Scutellastra) granatina Lam., Pilsbry, Man.
Conch., vol. 13, p. 106, pl. 62, figs. 76, 77.
1891 Patellona granatina Lam., Thiele, Das Gebiss der
Schnecken, vol. 2, p. 317.
1931 Patellona granatina Lam., Tomlin, Ann. Natal Mus.,
vol. 6 (3), p. 417 (designated type of Patellona)
1949 Patella granatina Lam., Koch, Ann. Natal Mus., vol. 11
(3), p. 501, pl. 20, figs. 1-5; text figs. 9, 10 (radula)
Records—SOUTH AFRICA: Table Bay (AWBP coll.); False
Bay (AWBP coll.); Platboom, Cape Peninsula (V. Orr, 1955;
ANSP); Sea Point (Mrs. N. Prior) Simonstown (AWBP coll.).
Patella oculus Born, 1778
(PI. 64, figs. 7-9; pl. 81; pl. 82, fig. 1)
Range—South Africa, west coast from near
Cape Town eastward to Umhlali.
Remarks—This large, depressed, broadly-ovate,
star-shaped limpet has something of the appear-
ance of Patella (Scutellastra) longicosta Lam-
arck, but differs from it, not only in dentition,
but also in coloration, for the interior of oculus is
dark purplish brown, except for a yellowish brown
spatula and a surrounding area of light bluish grey.
The species is essentially a cold-water one, and is
more abundant along the west coast of South
Africa than to the eastward. It occurs in the Bala-
noid zone, which is lower mid-tidal, but sometimes
extends to and below low spring-tide level.
Description—Shell large, up to 110 mm. (4%
inches) in length, solid, depressed, broadly ovate,
star-shaped, with the principal ribs strongly cor-
rugating the margin. Sculpture consisting of
about 11 primary, broad, carinated radials, plus
secondary radials and interstitial threads. Colour
of exterior dull-brown to blackish, but usually
eroded to dull-light greyish brown; interior with
a very broad dark purplish brown border, and a
light bluish grey area surrounding the spatula,
which is fawn to deep yellowish brown.
Radula—Formula 3 + | + (2+1+2) + 1 + 3, very
similar to the radula of granatina, the central
teeth having the same chevronlike alignment.
Measurements (mm.)—
length width height
110.0 106.0 42.0 Cape of Good Hope
86.0 76.0 15.0 Port Elizabeth
73.0 63.0 15.0 Port Alfred
53.0 51.0 8.0 Still Bay
Plate 81. Patella (Patellona) oculus Born, 1778. Cape of Good
Hope, South Africa, 77 mm., AWBP coll. 200.
[01-614]
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 111
Plate 82. Fig. 1. Patella (Patellona) oculus Born, South Africa.
Radula, from Koch, 1949, p. 508, fig. 18. Fig. 2 Patella (Pa-
tellona) granatina Linnaeus, South Africa. Radula, from Koch,
1949, p. 502, fig. 10. Fig. 3. Patella (Patellona) canescens
Gmelin, St. Helena. Radula, from Thiele, 1891, pl. 28, fig. 7
(as plumbea Lamarck). Fig. 4. Patella (Patellona) adansonii
Dunker, “Chinchao” in error, probably West Africa. Radula,
from Thiele, 1891, pl. 28, fig. 8.
Synonymy—
1778 Patella oculus Born, Index Mus. Caes. Vind., p. 434.
1786 Patella oculus hirci Lightfoot, Cat. Portland Mus., p.
105.
1791 Patella badia Gmelin, Syst. Nat., ed. 13, p. 3700.
1791 Patella monopis Gmelin, Syst. Nat., ed. 13, p. 3707.
1791 Patella fuscescens Gmelin, Syst. Nat., ed. 13, p. 3701.
1798 Patella astrolepas Réding, Mus. Bolten., vol. 2, p. 12.
1819 Patella scutellaris Lamarck, Anim. sans vert., vol. 6 (1),
p. 328.
1848 Patella schroeteri Krauss, Siidafr. Moll., Stuttgart, p.
43.
1854 Patella oculus Born, Reeve, Conch. Iconica, pl. 2, figs.
2a, b.
1891 Patella (Scutellastra) oculus Born, Pilsbry, Man. Conch.,
vol. 13, p. 106, pl. 27, figs. 30-32.
1932 Patella oculus Born, Turton, Mar. Shells Port Alfred,
p. 162.
1932 Patella oculus badia Gmelin, Turton, Mar. Shells Port
Alfred, p. 163.
1932 Patella oculus fuscescens Gmelin, Turton, Mar. Shells
Port Alfred, p. 163.
1932 Patella oculus schroeteri Krauss, Turton, Mar. Shells
Port Alfred, p. 163.
1932 Patella oculus planulata Turton, Mar. Shells Port Al-
fred, p. 163.
1942 Patella oculus Born, Tomlin & Stephenson, Proc. Malac.
Soc., Lond., vol. 25, pp. 5, 6.
1949 Patella oculus Born, Koch, Ann. Natal Mus., vol. 11 (3),
p. 507, pl. 22, figs. 1-4; text figs. 17, 18 (radula).
1967 Patella scutellaris Lamarck, Christiaens, Bull. Mus.
Nat. d’Hist. Nat. ser. 2, vol. 39 (5), p. 973.
Records—SOUTH AFRICA: west coast near Cape Town,
eastward to Umhlali (Koch, 1949, p. 507); Cape of Good Hope
(AWBP coll.);False Bay (AWBP coll.); Buffel’s Bay, Cape Pen-
insula (Mrs. N. Prior); Still Bay (Auck. Mus.); Port Elizabeth
(Auck. Mus.); Jeffrey's Bay (Auck. Mus.); Port Alfred (AWBP
coll.).
Patella adansonii Dunker, 1853
(Pl. 82, fig. 4; PI. 83, figs. 5, 6)
Range—West Africa.
Remarks—This species is characterised by its
dense fine radial ribbing, brown-lined and mar-
bled external pattern, and finely denticulated
margin.
Description—Shell of moderate size, 36-50 mm.
(1%-2 inches) in length, ovate, the anterior end
slightly narrowed, moderately elevated, with the
apex at about the anterior third. Sculpture con-
sisting of about 80 to 100 narrowly-rounded ra-
dial ribs that more or less alternate in strength.
Colour, externally whitish, marbled, and radially
and narrowly streaked with olive or dark greenish
brown, internally pale bluish grey, with the ex-
ternal pattern showing through, more strongly
at the margin; spatula buff to pale orange-brown.
Radula—Formula 3 + 1 + (2+1+2) + 1+ 3. The
five central teeth are not in a straight horizontal
row as in typical Patella, for the outer pair of
centrals are lower than the inner pair, and the
median one is small, very slender and almost
vestigial.
Measurements (mm.)—
length width height
50.0 41.0 17.0
42.0) 33.0 14.0
Pilsbry, 1891, p. 92
Angola
[01-615]
112. Subgenus Patellona A. W. B. Powell
Synonymy—
1853 Patella adansonii Dunker, Ind. Moll. Guin. Infer., p. 42,
pl. 6, figs. 10-15.
1891 Patella adansonii Dunker, Pilsbry, Man. Conch., vol. 13,
p. 92, pl. 12, figs. 30-33.
Records—WEST AFRICA: Loanda (type); Ambrizette, An-
gola (AWBP coll.; ANSP).
Patella canescens Gmelin, 1791
(Pl. 82, fig. 3; Pl. 83, figs. 3, 4)
Range—St. Helena.
Remarks—The sculpture is much finer than that
in either lugubris or plumbea and in conse-
quence the shell margin is delicately crenulated
rather than corrugated.
Plate 83. Figs. 1, 2. Patella (Patellona) lugubris Gmelin, 1791.
Porto Grande, St. Vincent, Cape Verde Islands, 42-54 mm.,
AWBP. coll. 50089. Figs. 3,4. Patella (Patellona) canescens
Gmelin, 1791. St. Helena, 41-45 mm., AWBP coll. 125412.
Figs. 5, 6. Patella (Patellona). adansonit Dunker, 1553,
Anvola, West Africa, 36-42 mim., AWBP coll. 146139.
Patellidae
Description—Shell of moderate size, up to 47
mm. (1% inches) in length, ovate, slightly nar-
rowed in front, moderately elevated, with the
apex subcentral. Sculpture crisp, consisting of
very numerous radial cords, the primaries
grouped in pairs or in threes, with an occasional
intermediate between each group of primaries.
Colour of exterior black, usually eroded to a
greyish brown. Interior bluish silvery, the spat-
ula flesh to orange-brown, and the edge of the
shell narrowly margined in black.
Measurements (mm.)—
length width height
47.0 40.0 20.0 St. Helena; Christiaens, 1968
45.0 37.5 20.0 St. Helena
Radula—F ormula 3 + 1 + (2+1+2) + 1 +3. Radula
very similar to that of plumbea and adansonii,
with the median central very small and the pairs
of centrals arranged chevron-like, descending
sharply from the central line. The lateral is dis-
tinctive in having four well developed cusps
(Christiaens, 1968, text fig. 1).
Synonymy—
1791 Patella canescens Gmelin, Syst. Nat., ed. 13, p. 3724.
Locality?
1855 Patella canescens Gmelin, Reeve, Conch. Iconica, pl. 34,
figs. 103 a, b. Locality?
1968 Patella canescens Gmelin, Christiaens, Rev. Zool. Bot.
Afr., vol. 77, pts. 3-4, pp. 314-320. St. Helena.
Patella lugubris Gmelin, 1791
(PI. 83, figs. 1, 2)
Range—West Africa, Loanda, Benguela, Guinea
and Cape Verde Islands.
Remarks—The species is much more coarsely
ribbed than either the St. Helena canescens or
the West African plumbea, and from the latter
species it differs in being more broadly ovate.
Description—Shell moderately large, up to 60
mm. (2% inches) in length, broadly ovate, mod-
erately elevated, with the apex almost at the
anterior third. Sculpture consisting of numerous
strong, keeled radials that prominently corrugate
the margin. Colour, externally dull-black, inter-
nally silvery bluish grey, the spatula often
clouded with a white callus.
Measurements (mm.)—
length width height
60.0 50.0 20.0 Pilsbry, 1891, p. 91
53.0 46.0 19.0 Cape Verde Islands
[01-616]
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, No. 15
Patella 113
Synonymy—
1791 Patella lugubris Gmelin, Syst. Nat. ed. 13, p. 3705;
based upon Martini-Chemnitz, Conch. Cab., vol. 1,
pl. 8, fig. 60.
1854 Patella lugubris Gmelin, Reeve, Conch. Iconica, pl. 14,
figs. 32 a-c. “Island of St. Vincent, West Indies,” sic.
- St. Vincent, Cape Verde Islands.
1891 Patella lugubris Gmelin, Pilsbry, Man. Conch., vol. 13,
p. 90, pl. 12, figs. 39, 40, 41-44; pl. 57, figs. 32-35.
Patella plumbea Lamarck, 1819
(PI. 84)
Range—Senegal, West Africa.
Remarks—This species is closely allied to lugu-
bris, and when adequate material is studied, may
prove to be identical. Pilsbry (1. c.) remarked
that “the ribbing is finer than in P. lugubris, the
shell is more elliptical, more depressed, and the
central spatula of the interior is longer and nar-
rower.”
Description—Shell moderately large, up to 53
mm. (2-1/16 inches) in length, narrowly ovate,
low-conical. Colour, externally dull-black, inter-
nally bluish, the spatula whitish, often clouded
with brown.
Measurements (mm.)—
length width height
52.0 38.0 10.5. type; Mermod, 1950, p. 692
Synonymy—
1819 Patella plumbea Lamarck, Anim. s. Vert., vol. 6, p. 328.
1834 Patella caerulea Quoy and Gaimard, Voy. Astrolabe,
Moll., vol. 3, p. 342, pl. 70, figs. 4-6.
1854 Patella plumbea Lam., Reeve, Conch. Iconica, pl. 3,
figs. 3a, b.
Plate 84. Patella (Patellona) plumbea Lamarck, 1819. Senegal,
West Africa. Holotype, 52 mm., from Mermod, Rev. Suisse
Zool., vol. 57, no. 34, p. 693, fig. 4.
1891 Patella plumbea Lam., Pilsbry, Man. Conch., vol. 13,
p. 91, pl. 24, figs. 11, 14, 15; pl. 57, figs. 38, 39.
1950 Patella plumbea Lam., Mermod. Rev. Suisse Zool., vol.
57, no. 34, pp. 692, 693, text fig. 4 (type).
Patella safiana Lamarck, 1819
(PI. 72, figs. 3, 4)
Range—Algeria, Morocco and West Africa.
Remarks—This is a large, elongate-ovate spe-
cies, only moderately elevated, and with a long
narrow spatula. In coloration the exterior is whit-
ish, with conspicuous brown rays in the rib inter-
stices. The interior is silvery grey, with the brown
external rays showing through towards the mar-
gin; the spatula is creamy-white, often stained
with orange-brown.
Description—Shell large, up to 77 mm. (3
inches) in length, elongate-ovate, moderately ele-
vated, with the apex at about the anterior third.
Sculptured with broadly rounded primary radial
ribs and weak interstitial cords. Colour: exter-
nally with whitish primary ribs and the interstices
intermittently rayed with brown; internally buff
to silvery-grey, slightly iridescent; spatula cream,
clouded with light orange-brown.
Radula—F ormula 3 + 1 + 4 + 1 + 3. The radula
resembles that of adansonii, canescens and plum-
bea, in that the centrals are not in a horizontal
line, the outer pair being lower than the inner
pair. Also, a median central appears to be com-
pletely absent, as in vulgata (see Fischer-Piette,
1935, p. 53, text fig. 22).
Measurements (mm.)—
length width height
77.0 57.0 21.0 largest of Lamarck’s type series
65.0 51.0 21.0 Oran, Algeria
Synonymy—
1819 Patella safiana Lamarck, Anim. sans vert., vol. 6, p. 329.
1849 Patella conspicua Philippi, Abbild., vol. 3, p. 71. Guinea.
1852 Patella kraussii Dunker, Index Moll. Guin. inf., p. 42,
pl. 6, figs. 4-6.
1854 Patella conspicua Philippi, Reeve, Conch. Iconica, pl. 7,
fig. 12. Gaboon.
1891 Patella safiana Lam. Pilsbry, Man. Conch., vol. 13, p. 90,
pl. 55, figs. 19-21.
1935 Patella safiana Lam. Fischer-Piette, Journ. Conchyl.,
vol. 79, p. 53.
1950 Patella safiana Lam., Mermod, Rev. Suisse Zool., vol. 57,
no. 34, pp. 693, 694, text fig. 5.
Types—The type series of safiana is in the Mu-
séum D’Histoire Naturelle de Geneve.
Records—ALGERIA: Oran (AWBP coll.). MOROCCO: ocean
coast (Pilsbry, 1891). WEST AFRICA: Gaboon, Guinea (Phi-
lippi, 1549, type of conspicua).
[01-617]
A. W. B. Powell Patellidae
114. Subgenus Patellona
[These occasional blank areas occur between
genera and subgenera to permit the insertion
of new material and future sections in their
proper systematic sequence. |
[01-618]
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 115
Subgenus Patellidea Thiele in Troschel, 1891
Type: Patella granularis Linnaeus, 1758
This subgenus, of which the type species is the
only known member, appears to be most closely
allied to the subgenus Scutellastra. The radula
resembles that of Scutellastra in its main features,
especially in having a narrow but well-developed
median central, and the other 4 larger centrals in
a horizontal line, but differs in the form of the
cusps which are oblique and parrot-beaklike.
The shell also differs from that of Scutellastra
in texture in being more strongly coloured, and in
having distinctive external sculpture, consisting
of strong radials bearing prominent imbricated
scales. Recent, South Africa.
Synonymy—
1891 Patellidea Thiele in Troschel, Das Gebiss der Schnecken,
volume 2, p. 315. Type, by monotypy: Patella granu-
laris Linnaeus, 1758.
Patella granularis Linnaeus, 1758
(pl. 64, figs. 4-6; pls. 85-87)
Range—South Africa, the entire coastline from
Port Nolloth in the west to Umpangazi in the east.
Remarks—This common South African species
is easily identified by its scaly external ribbing and
bluish white interior, broadly margined in dark-
brown, and with a reddish brown spatula.
Reeve’s Patella vidua, erroneously recorded
from the Philippines, is a synonym. A photograph
of one of Reeve’s figured specimens (Fig. 22a)
was kindly supplied by Dr. J. D. Taylor, and that
specimen, in the collections of the British Museum
(Natural History), is here nominated lectotype of
vidua.
Description—Shell of moderate size, up to 63
mm. (2% inches) in length, ovate, slightly narrowed
in front, tall-conical, with the apex slightly anter-
ior of the centre. Sculpture of strong, regular,
rounded primary radial ribs, with slightly weaker
intermediates; 1 to 3 intermediates between the
primary radials; all ribs with closely-spaced, im-
bricated, scales, resultant from numerous, lamel-
lose concentric growth lines. Colour: externally
dull light-brown to grey, with the scales paler;
internally bluish white, with a dark-brown, wide,
marginal border, and a reddish brown spatula. In
fully grown examples the border is usually contin-
uous, but in young shells it is interrupted by
bluish white radial streaks, corresponding with
the external radials.
Radula—Formula 3 + 1 + (2+1+2) + 1+ 3. Central
teeth 5, the middle member small and slender,
almost vestigial, the outer pairs much larger, each
with a long, pointed and incurved cusp; pluri-
cuspid lateral with an enlarged top, bearing 4
cusps, of which the second from the proximal side
is largest, and shaped like those of the central
pairs; marginals 3, long, narrow and _ flexuous,
each with a rudimentary cusp.
Plate 85. Patella (Patellidea) granularis Linnaeus, 1758. Figs.
1, 2. Platboom, Cape Point, South Africa. Fig. 3. Cape of
Good Hope, South Africa, 37-60 mm., AWBP coll. 42924 &
193.
[01-629]
116 Subgenus Patellidae
Measurements (mm.)—
length width height
63.0 48.0 19.0 Sea Point; Mrs. N. Prior
59.0 49.0 26.0 C. of Good Hope
49.0 38.0 24.0 Platboom
44.5 36.0 17.0 Port Alfred
Synonymy—
1758 Pateklla granularis Linnaeus, Syst. Nat., ed. 10, p. 782.
1834 Patella granularis L., Quoy and Gaimard, Voy. ‘Astro-
labe’, Zool. vol. 3, p. 341, pl. 70, figs. 12-15.
1848 Patella. granularis’ L., Krauss, Sudafr. Moll., Stuttgart,
p. 52.
1848 Patella echinulata Krauss, Sudafr. Moll., Stuttgart, p. 52,
pl. 3, fig. 15.
1848 Patella natalensis Krauss, Sudafr. Moll., Stuttgart, p. 53,
pl. 3, fig. 10.
1854 Patella vidua Reeve, Conch. Iconica, pl. 11, figs. 22a, b.
1854 Patella granularis L., Reeve, Conch. Iconica, pl. 14, figs.
31a, b.
1855 Patella morbida Reeve, Conch. Iconica, pl. 25, figs. 64a,
b.
1891 Patella (Scutellastra) granularis L., Pilsbry, Man. Conch.,
vol. 13, p. 102, pl. 63, figs. 80-83.
1891 Patellidea granularis L., Thiele (new genus), in Troschel
& Thiele, Das Gebiss der Schnecken, col. 2, p. 315.
1931 Patellidea granularis Linne, (designated type of genus)
Tomlin, Ann. Natal. Mus., vol. 6 (3), p. 417.
1932 Patella granularis L., Turton, Mar. Shells Port Alfred,
p. 166.
Plate 86. Patella (Patellidea) granularis Linnaeus, 1758. Lec-
totype, here nominated, of Patella vidua Reeve, 1854, er-
roneously recorded from the Philippines, but considered to
be synonymous with the South African granularis, The lecto-
type is based upon Reeve’s fig. 22a. Photo by courtesy of
Dr. J. D. Taylor, British Museum (Natural History).
A. W. B. Powell
Patellidae
1932 Patella morbida Reeve, Turton, Mar. Shells Port Alfred,
p. 166.
1932 Patella natalensis Krauss, Turton, Mar. Shells Port
Alfred, p. 166.
1932 Patella natalensis echinulata Krauss, Turton, Mar. Shells
Port Alfred, p. 166.
1932 Patella miliaris Turton, Mar. Shells Port Alfred, p. 166.
(non Philippi, 1848)
1932 Patella argenvillei assimilans Turton, Mar. Shells Port
Alfred, p. 167.
1932 Patella alboradiata Turton, Mar. Shells Port Alfred,
p. 167.
1933 Patella tomlini Turton, (nom. nov. pro P. alboradiata
Turton, 1932, non Gmelin, 1791) Journ. Conch., vol.
19, p. 371.
1949 Patella granularis Linne Koch, Ann. Natal Mus., vol. II
(3), p. 503, pl. 19, figs. 4-8; text figs. 11,12 (radula).
Types—The types of granularis (Holotype;
Sloane coll., no. 1013), morbida and vidua (lecto-
type, here selected) are in the Britsh Museum
(Natural History).
Records—SOUTH AFRICA: Port Nolloth to Umpangazi,
north of Durban (Koch, 1949, p. 503); Saldanha Bay (Dis-
covery II, 1926); Table Bay (AWBP coll.); Cape of Good Hope
(AWBP coll. 193); Sea Point (Mrs. N. Prior); False Bay (Auck.
Mus.); Platboom, Cape Point (V. Orr, Jan. 1955); Jeffrey's
Bay (AWBP coll.); Port Alfred (Auck. Mus.).
K,
|
Plate 87. Fig. 1. Patella (Patellidea) granularis Linnaeus.
South Africa. Radula. Fig. 2. Patella (Olana) cochlear Born.
Radula, both from Koch, 1949, p. 504, fig. 12 & p. 499,
fig. 6.
[01-630]
November 27, 1973
Subgenus Cymbula H. and A. Adams, 1854
Type (monotypy): Patella compressa Linnaeus, 1758
This subgenus contains two species, the shells
of which are of very different outward appearance,
but nevertheless have a striking sameness in the
radula that is of a distinctive type. The multiple
centrals and the pluricuspid lateral have large
blunt-topped cusps, with strongly raised or
flanged edges. The centrals collectively form a
chevron instead of forming a horizontal line, as in
typical Patella, and thus indicate alliance with the
subgenus Patellona.
The type species is the easily recognised com-
pressa, with its elongated, laterally compressed
shell, adapted to its specialised station on the
stipes of the large kelps, Ecklonia and Laminaria.
Only rarely is this species found attached to rock.
On the other hand, the second species, miniata,
is of normal ovate limpet shape, since it is ex-
clusively a rock-dwelling. Both species are re-
stricted to South African waters.
Synonymy—
1854 Cymbula H. Adams and A. Adams, The Genera of Re-
cent Mollusca, volume 1, p. 466. Type, by monotypy:
Patella compressa Linnaeus, 1758. [Cymbula Gray,
1821, is an error for Cymbulia Peron and Lesnerr
1810].
Patella compressa Linnaeus, 1758
(pl. 63, fig. 9; Pls. 88, 89)
Range—South Africa, from Port Nolloth in the
west to Danger Point in the south. Records from
further afield, including one from St. Helena, are
due to drift, along with large algae, upon which
the species lives.
Description—Shell large, up to 117.5 mm. (4%
inches) in length, thin, elongate-ovate, tall and
narrow, with parallel sides, the apex a little for-
ward of the middle, and curving anteriorly. Sculp-
ture consisting of very numerous, rather unequal,
linear-spaced riblets; margin very minutely crenu-
lated, convex at the sides, and concave at the
ends. Colour: externally dull brownish buff; inter-
nally light pinkish fawn, the central area irregu-
larly clouded with whitish callus.
Radula—F ormula 3 + 1 + (2+1+2) + 1 + 3. Cen-
tral teeth 5, forming a chevron, the median one
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patella 117
small and very narrow, bearing a simple small
cusp, outer pairs of centrals massive, each with a
broa,d flat-topped cusp, ridged on each side, the
ridges more prominent on the outermost teeth;
pluricuspid lateral with two cusps, each similar to
those of the outer centrals; the three marginals
are small, each with a single simple cusp, outer-
most marginal largest of the three.
Measurements (mm.)—
length width height
117.5 51.0 50.0 Kommetje, Cape Peninsula;
Mrs. N. Prior.
94.0 45.0 35.0 South Africa; Pilsbry, 1891,
p. 93
83.0 44.0 36.5 South Africa
63.5 35.0 23.0 South Africa
Synonymy—
1758 Patella compressa Linnaeus, Syst. Nat. ed. 10, p. 783.
1834 Patella compressa L., Quoy and Gaimard, Voy. ‘Astro-
labe’, Zool., vol. 3, p. 338, pl. 70, figs. 1-3.
1848 Patella compressa L., Krauss, Siidafr. Moll., Stuttgart,
p. 50.
1854 Patella compressa L., Reeve, Conch. Iconica, pl. 7, figs.
13a, b.
1854 Patella (Cymbula) compressa L., H. & A. Adams, Gen.
Rec. Moll., vol. 1, p. 466.
1891 Patella compressa L., Pilsbry, Man. Conch., vol. 13, p. 93,
pl. 61, figs. 68-70.
1949 Patella compressa Linne. Koch, Ann. Natal Mus., vol. 11,
p. 499, pl. 17, figs. 4-6; text figs. 7,8 (radula).
Records—SOUTH AFRICA: Port Nolloth to Danger Point
(Koch, 1949, p. 499); Cape Peninsula (AWBP coll. 26039);
Kommetje, Cape Peninsula (Mrs. N. Prior).
Plate $8. Fig. 1. Patella (Cymbula) compressa Linnaeus.
South Africa. Radula, from Koch, 1949, p. 500, fig. 8. Fig. 2.
Patella (Cymbula) miniata Born. South Africa. Radula, from
Koch, 1949, p. 507, fig. 16.
[01-635]
118 = Subgenus Cymbula
Patella miniata Born, 1778
(Pl. 63, figs. 10, 11; Pls. 88, 90, 91)
Range—South Africa, from Port Nolloth in the
west, eastward to Qolora, near East London, and
Natal.
Remarks—This moderately large, very attractive
limpet is ovate, depressed and relatively thin,
with an intricate pattern of radial streaks and
speckles; it is reddish brown in living examples,
but bleached to bright-pink in shells from beach
drift. Pilsbry (1891, p. 93) was incorrect in as-
suming that miniata is merely a rock-dwelling
ecotype of the kelp living compressa.
Description—Shell rather large, up to 93 mm.
3% inches) in length, strong but relatively thin,
ovate, slightly attenuated in front, rather de-
pressed, and with the apex varying between sub-
A. W. B. Powell
Patellidae
central and the anterior third. Sculpture consist-
ing of numerous primary radial cords, with
mostly two radial threads in the interspaces; the
ribbing varies in strength, and may be almost
smooth to sharply-imbricated by dense concen-
tric growth threads. Colour: externally radially
streaked and speckled in reddish brown to bright-
pink, on a white ground; internally silvery pinkish
white, with the external pattern showing through
strongly; spatula white-callused, sometimes tinged
with orange. Living examples are usually en-
crusted.
Radula—Formula 3 + 1 + (2+1+2) + 1 +3. The
radula stands nearest to that of compressa, the
arrangement of the teeth being the same. The
only noticeable difference between the two is in
the shape of the cusps which have convex cutting
edges in miniata but straight to concave ones in
compressa.
Plate 89. Patella (Cymbula) compressa Linnaeus, 1758.
South Africa, 64-83 mm., AWBP coll. 1403 & 17985.
[01-636]
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 119
Plate 90. Figs. 1-3. Patella (Cymbula) miniata Born, 1778,
South Africa. Figs. 1, 2. Port Nolloth, 75 mm., AWBP. coll.
Measurements (mm.)—
length width height
93.0 71.5 22.0 Natal; Mrs. N. Prior.
79.0 66.0 20.0 False Bay
75.0 56.5 16.0 Port Nolloth
54.0 43.5 12.0 Still Bay
Synonymy
1778 Patella miniata Born, Index Mus. Caes. Vind., p. 436;
1780, Test. Mus. Caes. Vind., p. 420.
1786 Patella pulchra Lightfoot, Cat. Portland Mus., p. 105.
1791 Patella umbella Gmelin, Syst. Nat., ed. 13, p. 3706.
1791 Patella sanguinolenta Gmelin, Syst. Nat., ed. 13, p. 3716.
1798 Patella rosea Réding, Mus. Bolten., vol. 2, p. 9.
1798 Patella rubicunda Réding, Mus. Bolten, vol. 2, p. 9.
1848 Patella miniata Born, Krauss, Sudafr. Moll., Stuttgart,
p. 51.
1854 Patella sanguinans Reeve, Conch. Iconica, pl. 6, fig. 10.
1854 Patella umbella Gmelin, Reeve, Conch. Iconica, pl. 9,
figs. 17a, b.
1891 Patella. compressa var miniata Born, Pilsbry, Man.
Conch., vol. 13, p. 94, pl. 26, figs. 22-27.
1932 Patella miniata Born, Turton, Mar. Shells Port Alfred,
p. 168.
1932 Patella miniata umbella Gmelin, Turton, Mar. Shells
Port Alfred, p. 168.
1932 Patella miniata decorata Turton, Mar. Shells Port Alfred,
p. 169. pl. 39, fig. 1199.
1932 Patella pulchella Turton, Mar. Shells Port Alfred, p. 169,
pl. 39, fig. 1200. (non Blainville, 1825).
1932 Patella alboguttata Turton, Mar. Shells Port Alfred,
p. 169, pl. 39, fig. 1202.
1932 Patella denseplicata Turton, Mar. Shells Port Alfred,
p. 169, pl. 39, fig. 1205.
1932 Patella densestriata Turton, Mar. Shells Port Alfred,
p. 170, pl. 39, fig. 1206.
1933 Patella becki Turton, Journ. Conch., vol. 19, p. 371;
nom. nov. pro P. pulchella Turton, 1932, non Blain-
ville, 1825,
52455. Fig. 3. False Bay, 79 mm., AWBP coll. 26041.
1942 Patella sanguinolente (sic Gmelin, Tomlin & Stephenson,
Proc. Malac. Soc., Lond., vol. 25, p. 7.
1949 Patella miniata Born, Koch, Ann. Natal Mus., vol. 11
(3), p. 506, pl. 21, figs. 1-12; text figs. 15a, b, 16
(radula).
Records—SOUTH AFRICA: Port Nolloth in the west to
Qolora in the east, common in the sub-littoral fringe (Koch,
1949, p. 506); Port Nolloth; False Bay; Still Bay; Port Elizabeth;
Algoa Bay (all AWBP coll.); Natal (Mrs. N. Prior).
Plate 91. Patella (Cymbula) miniata Born, 1778. Natal,
South Africa; An extra large and fine example of the species,
in the collection of Mrs. Nancy Prior of Cape Town. It has a
length of 93 mm. (3% inches).
[01-637]
120
Subgenus Cymbula
A. W. B. Powell
Patellidae
[These occasional blank areas occur between
genera and subgenera to permit the insertion
of new material and future sections in their
proper systematic sequence. |
[01-635]
November 27, 1973
Subgenus Olana H. and A. Adams, 1854
Type (monotypy): Patella cochlear Born, 1778
A moderate-sized shell of depressed pear-
shape, with the anterior end laterally constricted
and produced like a spout. The sole species of
this subgenus is restricted to South Africa where
in many places it is so abundant that it forms a
dense mosaic, termed the “Cochlea zone.” A
density of 1,300 examples to the square-yard
has been recorded, and as many as 40 crowded
on top of a single large shell. Almost all large
examples bear several deeply excavated scars,
resultant from superimposed individuals. These
limpets do not appear to move around much, but
merely rotate, so that the head can move in a
circle and the radula crop the algal growth within
its range (See Koch, 1949, pp. 498-499).
Synonymy—
1854 Olana H. and A. Adams, The Genera of Recent Mol-
lusca, vol. 1, p. 466. Type by monotypy: Patella coch-
lear Gmelin, 1791 = Born, 1778.
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella 121
Patella cochlear Born, 1778
(Pl. 63, figs. 12-14; Pls. 87, 92)
Range—South Africa, from Buffalo River on the
west coast and around the Cape of Good Hope to
Port Edward, Natal.
Description—Shell of moderately large size, up
to 67 mm. (2% inches) in length, solid, depressed,
pear-shaped, with the anterior end much con-
stricted, and produced like a spout. Sculptured
with strong, rather unequal radial ridges that
deeply corrugate the margin. Colour: externally
white to yellowish brown; internally white, tinged
with bluish grey; the spatula fawn, orange-brown,
or clouded with white, and surrounded, except in
front of the head region, with a broad band of
indigo.
Radula—Formula 3 + 1 + (2+1+2) + 1 + 3. The
small, slender, median central tooth, and the two
pairs of multiple centrals form a horizontal line,
as in true Patella, but the massive lateral has
four cusps, two of them large and the other two
much smaller. The cusps are heart-shaped, with
a pronounced groove down the middle; marginals
3, each with a weak cusp.
Measurements (mm.)—(all A. W. B. Powell col-
lection).
length width height
67.0 50.0 17.0 Port Alfred
58.5 43.5 15.5 Cape Natal
45.0 34.5 6.0 False Bay
Plate 92. Patella (Olana) cochlea Born, 1778, Port Alfred,
South Africa, 66 mm., AWBP coll. 51974.
[01-643]
122. Subgenus Olana
A. W. B. Powell
Patellidae
Synonymy—
1778 Patella cochlear Born, Index Mus. Caes. Vind., p. 437;
1780, Test. Mus. Caes. Vind., pl. 18, fig. 3, p. 420.
1790 Patella cochlear Born, Gmelin, Syst. Nat. ed. 13,
p. 3721.
1848 Patella cochlear Born, Krauss, Sudafr. Moll., Stuttgart,
p. 48.
1854 Patella cochlear Born, Reeve, Conch. Iconica, pl. 12,
figs. 24a, b.
[01-644]
1854 Patella (Olana) cochlear Born, H. & A. Adams, Gen.
Rec. Moll., vol. 1, p. 466.
1891 Patella (Scutellastra) cochlear Born, Pilsbry, Man.
Conch., vol. 13, p. 104, pl. 27, figs. 34, 35.
1949 Patella cochlear Born, Koch, Ann. Natal Mus., vol. 11
(3), p. 498, pl. 19, figs. 1-3; text figs. 5, 6 (radula).
Records—SOUTH AFRICA: False Bay; Sea Point (Mrs. N.
Prior); Port Elizabeth; Cape Natal; Port Alfred (all AWBP
coll.).
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patella = 123
nn.
Subgenus Scutellastra H. and A. Adams, 1854
Type: Patella barbara Linnaeus, 1755
Mostly large massive shells with an opaque,
non-iridescent, porcellaneous interior. The exter-
nal coloration is either a uniform spread of
colour, or the pigment may be confined to the
spaces between the radial ribs and show through
to the inside margin to form a narrow border
where the shell is thinnest.
Radula and gill cordon as in typical Patella,
but the median central tooth is usually well-de-
veloped, often as large as the other four centrals.
The range of the subgenus is South Africa, on
across the Indo-Pacific as far east as the Society
Islands, northward to Japan and along the south
coast of Australia. Undoubted fossil occurrences
of the subgenus are cooperi (Powell, 1935) from
the lower Miocene of Motuihi Island, Auckland,
New Zealand, and aurorae Fleming, 1973, from
the middle Oligocene of Mason River, North
Canterbury, New Zealand.
Synonymy—
1854 Scutellastra H. Adams and A. Adams, The Genera of
Recent Mollusca, volume 1, p. 466, for gorgonica
Hymphrey, pentagona Born and plicata Born. Type,
by subsequent designation, Powell, 1938: Patella gor-
gonica Humphrey “=P. longicosta Lamarck” (sic) =Pa-
tella barbara Linnaeus, 1758.
1924 Patellanax Iredale, Proceedings of the Linnean Society
of New South Wales, volume 49, part 3, p. 239. Type,
by original designation: Patella squamifera Reeve,
1855.
1929 Penepatella Iredale, Memoirs of the Queensland Mu-
seum, volume 9, part 3, p. 276. Type, by original
designation: Penepatella inquisitor Iredale, 1929.
Patella argenvillei Krauss, 1848
(PI. 65, fig. 4; Pls. 93, 96)
Range—South Africa: Port Nolloth on the west
coast, eastward to Qolora, between East London
and Durban.
Remarks—This species is more common on the
west coast where it forms concentrated low-tidal
communities, termed the Cochlear-argenvillei zone.
It is one of the least variable of South African lim-
pets, easily recognised by its narrowly elongate-
oval, high-conical form, dense regular sculpture,
and dark external coloration, with white linear
rib interstices.
Description—Shell large, up to 89 mm. (34 inches)
in length, rather solid, oblong-ovate, slightly con-
stricted at the anterior end, high conical, with the
apex subcentral, a little nearer to the anterior end.
Sculpture consisting of very numerous, over 100,
more or less regular, flat-topped radial ribs, with
linear interstices; margin delicately and evenly cren-
ulated. Colour: externally blackish, the rib inter-
stices white; internally white, dark-greyish between
the marginal crenulations, and spatula diffused with
yellowish brown.
Radula—F ormula 3 + 1 + (2+1+2) + 1+ 3. Central
teeth consisting of a narrow insignificant middle
member, with a pair of strong, sharply-cusped and
centrally-grooved teeth on each side; lateral massive,
with an expanded top, bearing four sharp cusps,
the middle two deeply grooved; marginals three,
weakly cusped and slender.
Measurements (mm.)—
length width height
89.0 68.0 52.0 Table Bay
rae) 58.0 35.5 South Africa
63.0 44.5 30.0 Port Elizabeth
Plate 93. Patella (Scutellastra) argenvillet Krauss, 1545,
Port Alfred, South Africa, 72-77 mim., AWBP coll. 30065.
[01-649]
124. Subgenus Scutellastra
Synonymy—
1848 Patella argenvillei Krauss, Sudafr. Moll., Stuttgart, p. 49;
based upon Argenville Conch., 1870, vol. 1, p. 504 and
vol. 3, pl. 3, fig. G.
1854 Patella argenvillii Krauss, Reeve, Conch. Iconica, pl. 10,
figs. 20a, b.
1891 Patella (Scutellastra) argenvillei Krauss, Pilsbry, Man.
Conch., vol. 13, p. 95, pl. 22, figs. 15, 16; pl. 58, fig. 44.
1949 Patella argenvillei Krauss, Koch, Ann. Natal Mus., vol.
11(3), p. 494, pl. 17, figs. 1-3; text figs. 1, 2 (radula).
Records—SOUTH AFRICA: Table Bay (AWBP coll.); False
Bay (AWBP coll.); Sea Point (Mrs. N. Prior); Port Elizabeth
(Auck. Mus.); Port Alfred (AWBP coll.).
Patella barbara Linnaeus, 1758
(Pl. 65, figs. 1-3; Pls. 94-96)
Range—South Africa, the whole length of the
coastline from Port Nolloth in the west to Umpan-
gazi in the east.
Remarks—This is a large, solid, ovate limpet,
varying greatly in height, but always strongly
radially ridged, resulting in a deeply corrugated
margin. The coloration is buff to pale brownish
externally and white within, except for the spatula,
which is often blotched with reddish brown.
Description—Shell rather large, up to 95 mm. (3%
inches) in length, of only moderate height and
weight in its younger stages but tall and crass in
mature examples, narrowly to broadly ovate, with
the apex near central. Sculpture variable, but always
strongly and coarsely radially ribbed, their terminal
points corrugating the margin, sometimes almost as
Plate 94 Patella (Scutellastra) barbara Linnaeus, 1758, Port
A. W. B. Powell
Patellidae
strongly as in longicosta. Radial ribs carinated and
of varying strength, from 10 to 20 primaries and 1 to
4 secondaries in the interspaces, the whole rendered
noticeably scabrous by close set, lamellose, concen-
tric growth marks. The posterior end of the shell
usually has 5 ribs much stronger than the rest.
Colour: externally dull-buff to light yellowish
brown; internally whitish, often with a narrow,
pale-fawn, marginal border; spatula either irregu-
larly blotched with reddish-brown, or callused
over with white. A variable species, as shown by
the lengthy synonymy.
Radula—F ormula 3+ 1 + (2+1+2) + 1+3, similar to
that of both longicosta and cochlear in the form of
the massive lateral, which has an expanded head,
bearing three cusps, the middle one much the larger,
and there is an incipient fourth cusp on the outer
side. The five centrals have the middle member
small and slender, with a minute vestigial cusp,
but the cusps of the outer pairs of centrals, and
the middle member of the laterals have broad flat
tops, with ridged margins; marginals 3, slender,
flexuous, and each with a weak cusp.
Measurements (mm.)—
length width height
95.0 81.0 37.0 Buluga Bay, East London;
Mrs. N. Prior
95.0 70.0 31.0 var. ovalis Pilsbry, 1891, p. 97
84.5 61.5 31.5 Still Bay
79.0 58.0 38.0 CC. of Good Hope
79.0 62.0 22.5 Port Alfred
72.0 60.0 27.0 Pilsbry, 1891, p. 96
Alfred, South Africa, 76 mm., AWBP coll. 30063.
[01-650]
November 27, 1973
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patella = 125
Synonymy—
1758 Patella barbara Linnaeus, Syst. Nat., ed. 10, p. 782.
1778 Patella plicata Born, Mus. Caes. Vind., p. 433; 1780, Test.
Mus. Caes. Vind., p. 417, pl. 18, fig. 1.
1786 Patella fungoides Lightfoot, Cat. Portland Mus., p. 55.
1786 Patella gorgonica Lightfoot, Cat. Portland Mus., p. 105.
1791 Patella plicaria Gmelin, Syst. Nat., ed. 13, p. 3708.
1791 Patella cypria Gmelin, Syst. Nat., ed. 13, p. 3698.
1819 Patella barbata Lamarck, Anim. sans vert., vol. 6 (1),
p. 326.
1819 Patella spinifera Lamarck, Anim. sans vert., vol. 6 (1),
p. 326.
1848 Patella barbara L., Krauss, Sudafr. Moll., Stuttgart, p. 45.
1848 Patellaa obtecta Krauss, Sudafr. Moll., Stuttgart, p. 47,
pl..3, fig. 11.
1854 a le Born, Reeve, Conch. Iconica, pl. 9, figs.
l6a, b.
1891 Patella (Scutellastra) barbara L., Pilsbry, Man. Conch.,
vol. 13, p. 96, pl. 15, figs. 1, 2; pl. 59, figs. 50-55.
1591 Patella (Scutellastra) barbara var. ovalis Pilsbry, Man.
Conch., vol. 13, p. 97, pl. 60, figs. 56-58.
1932 Patella barbara L., Turton, Mar. Shells Port Alfred, p. 163,
sp. 1162.
1932 Patella barbara plicata Born, Turton, Mar. Shells Port Al-
fred, p. 163.
1932 Patella whitechurchi Turton, Mar. Shells Port Alfred, p.
164, pl. 37, fig. 1165.
1932 Patella thetis Turton, Mar. Shells Port Alfred, p. 164, pl.
37, fig. 1166.
1932 Patella hera Turton, Mar. Shells Port Alfred, p. 164, pl. 37,
fig. 1167.
1932 Patella amphitrite Turton, Mar. Shells Port Alfred, p. 164,
pl. 37, fig. 1168.
1932 Patella amphitrite var. brunescens Turton, Mar. Shells
Port Alfred, p. 164, pl. 37, fig. 1169.
1932 Patella decemcostata var. major Turton, Mar. Shells Port
Alfred, p. 165, pl. 35, fig. 1171.
1932 Patella nympha Turton, Mar. Shells Port Alfred, p. 165,
pl. 38, fig. 1174.
1932 Patella sowerbyi Turton, Mar. Shells Port Alfred, p. 166,
pl. 38, fig. 1176.
1949 Patella barbara Linne, Koch, Ann. Natal Mus., vol. 11 (3),
p. 496, pl. 18, figs. 1-12; text figs. 3a, b, 4 (radula).
Records SOUTH AFRICA: whole length of coastline from
west to east (Koch, 1949, p. 496); Table Bay (Auck. Mus.
3081); Cape of Good Hope; Still Bay; False Bay; Port Al-
fred; Cape Natal (all AWBP coll.).
Patella longicosta Lamarck, 1819
(Pl. 65, figs. 5-7; Pls. 95, 96)
Range—South Africa, from Oudekraal, west
side of Cape Peninsula, eastward to Umpangazi,
north of Durban.
Remarks—This species is easily recognised by its
depressed star-shape, with the primary ribs extend-
ing well-beyond the margin, which is narrowly bor-
dered in black. This species occurs commonly in the
lower Balanoid and Cochlear zones.
Description—Shell rather large, up to 76 mm. (3
inches) in length, solid, depressed, stellate, very
strongly sculptured with sharply carinated, radial
ridges, that project well-beyond the margin, seven
of them much stronger than the rest; apex at an-
terior third to submedian. Colour of exterior dull-
black, when not eroded, to a rusty-brown; interior
bluish white, with a narrow black margin, and a
yellowish brown spatula, the latter clouded with a
white callus in senile specimens.
Radula—F ormula 3+ 1 + (2+1+2) + 1+3, somewhat
similar to that of cochlear in the form of the cusps,
which are leaf-shaped with a median groove, and in
the massive lateral that has four cusps.
Plate 95. Patella (Scutellastra) longicosta) Lamarck, 1519.
Mossel Bay, Cape Peninsula, South Africa, 66 mm., AWBP
coll. 42429.
[01-651]
126 Subgenus Scutellastra A. W. B. Powell Patellidae
a Se ee
Measurements (mm.)— 1901 Patella decemcostata E. A. Smith, Journ. Conch., vol. 10,
. p. 106, pl. 1, fig. 22.
length width height 1932 Patella longicosta Lam., Turton, Mar. Shells Port Alfred,
75.5 75.5 23.0 False Bay p. 161, pl. 37, fig. 1145.
70.0 64.0 18.0 False Bay 1932 Patella longicosta intermedia Turton, Mar. Shells Port Al-
61.0 56.0 12.0 Cape Peninsula fred, p. 161, pl. 37, fig. 1146.
1932 Patella multilirata Turton, Mar. Shells Port Alfred, p. 161,
Synonymy— pl. 37, fig. 1147.
1932 Patella tabularis Krauss, Turton, Mar. Shells Port Alfred, p.
1819 Patella longicosta Lamarck, Anim. sans vert., vol. 6 (1), 161. (non Krauss, 1848; in part, smaller of two examples).
p. 326. 1932 Patella tabularis angulosa Gmelin, Turton, Mar. Shells
1842 Patella longicosta Lam., Reeve, Conch. Syst., vol. 2, p. 15, Port Alfred, p. 161, sp. 1150. (non Gmelin, 1791).
pl. 136, fig. 6. 1932 Patella tabularis monopsis Gmelin, Turton, Mar. Shells Port
1848 Patella longicosta Lam., Krauss, Sudafr. Moll., Stuttgart, Alfred, p. 162. (sic; non monopis Gmelin, 1792).
p. 48. 1932 Patella tabularis squamosa Gmelin, Turton, Mar. Shells
1854 Patella longicosta Lam., Reeve, Conch. Iconica, pl. 6, figs. Port Alfred, p. 161, sp. 1149. (non Gmelin, 1791).
lla, b. 1932 Patella granatina Linn. Turton, Mar. Shells Port Alfred,
1891 Patella (Scutellastra) longicosta Lamarck, Pilsbry, Man. p. 163, sp. 1161. (non Linnaeus, 1758).
Conch., vol. 13, p. 107, pl. 28, figs. 37, 38. 1932 Patella decemcostata E. A. Smith, Turton, Mar. Shells Port
Alfred, p. 165.
1932 Patella albanyana Turton, Mar. Shells Port Alfred, p. 165,
pl. 38, fig. 1175.
1933 Patella longicosta kowiensis Turton, Journ. Conch., vol. 19,
p. 371; nom. nov. pro P. longicosta intermedia Turton,
1932, non Knapp, 1857.
1942 Patella longicosta Lam., Tomlin and Stephenson, Proc.
Malac. Soc., Lond., vol. 25, pp. 4-9.
1949 Patella longicosta Lam., Koch, Ann. Natal Mus., vol. 11 (3),
p. 504, pl. 20, figs. 6-13; text figs. 13a-c, 14 (radula).
Records—SOUTH AFRICA: Cape Peninsula (AWBP coll.);
Kommetje, Cape Peninsula (Mrs. N. Prior); Mossel Bay, Cape
Peninsula (V. Orr, 1955; ANSP); False Bay (AWBP coll.); Still
Bay (Auck. Mus.); Simon’s Bay (V. Orr, 1955; ANSP); Port Eliza-
beth (Auck. Mus.); Cape Natal (AWBP coll.); Port Alfred (AWBP
coll.).
Types—The type series of three examples of longi-
costa is in the Muséum D’Histoire Naturelle de
Geneve.
Patella tabularis Krauss, 1848
(Pl. 66, fig. 5; Pls. 96, 97)
Range—South Africa, from Cape Peninsula east-
ward to Port St. John’s.
Remarks—This is the largest of the South African
limpets; it somewhat resembles kermadecensis but
has much more prominent radial sculpture. The
species inhabits the sub-littoral fringe, and does not
occur in dense communities (Koch, 1949, p. 509).
Description—Shell very large and massive, up to
147.5 mm. (5% inches) in length, broadly ovate, mod-
erately elevated, and with the apex anterior to the
middle. Sculpture heavy and coarse, consisting of
9, 10 or more heavy, foldlike, radial ribs, and nu-
merous secondary ribs of several sizes, the whole
imbricated by dense lamellose growth lines; margin
deeply and somewhat irregularly scalloped. Colour:
exterior dull reddish or rusty-brown; interior porcel-
Plate 96. Fig. 1. Radula of Patella (Scutellastra) barbara lanous-white, with a moderately wide border of
Linnaeus. Fig. 2. Patella (Scutellastra) longicosta Lamarck. ans : . ; ‘ =
Fig. 3. Patella (Seutellastra) argenvillei Krauss. Fig. 4. reddish brown, being the external colour show ane
Patella (Scutellastra) tabularis Krauss. South Africa. Radulae, through at the thinner margin; spatula not differen-
all from: Koch, 1949. tiated by colour.
[01-652]
November 27, 1973
Plate 97. Patella (Scutellastra) tabularis Krauss, 1848. Port
Alfred, South Africa, 115 mm., AWBP coll. 30062.
Measurements (mm.)—
length width height
147.5 125.5 50.0 Buluga Bay, East London;
Mrs. N. Prior
127.0 111.9 46.0 South Africa
114.0 98.0 32.0 Cape Point
81.0 68.0 20.5. Port Alfred
Radula—F ormula 3 + 1 + (2+1+2) + 1 + 3. Central
teeth consisting of a slender, almost vestigial, middle
member, flanked by pairs of large, blunt-cusped
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15 Patella = 127
teeth; lateral massive, with an expanded head, bear-
ing four cusps, the middle pair the stronger; mar-
ginals three, narrow and slender, each with a weak
cusp.
Synonymy—
1848 Patella tabularis Krauss, Sudafr. Moll., Stuttgart, p. 47,
pl: 3; fig. 8.
1854 Patella rustica L., Reeve, Conch. Iconica, pl. 5, figs. 8a, b.
(non Linnaeus, 1758).
1891 Patella (Scutellastra) tabularis Krauss, Pilsbry, Man.
Conch., vol. 13, p. 105, pl. 16, figs. 9, 10.
1891 Patella (Scutellastra) patriarcha Pilsbry, Man. Conch., vol.
13, p. 105, pl. 64, figs. 84, 85; pl. 65, fig. 86.
1932 Patella rustica L. Turton, Mar. Shells Port Alfred, p. 162,
sp. 1154.
1932 Patella tabularis Krauss, Turton, Mar. Shells Port Alfred,
p. 161, sp. 1148 (in part; larger of two specimens).
1932 Patella obtecta Krauss Turton, Mar. Shells Port Alfred,
p. 162, sp. 1155. (non Krauss).
1932 Patella patriarcha Pilsbry, Turton, Mar. Shells Port Alfred,
p. 162, sp. 1153.
1942 Patella tabularis Krauss Tomlin and Stephenson, Proc.
Malac. Soc., Lond. vol. 25 (1), p. 5.
1949 Patella tabularis Krauss, Koch, Ann. Natal Mus., vol. 11 (3),
p. 509, pl. 22, figs. 5-11; text figs. 19a, b, 20 (radula).
Types—The type of tabularis is in the Stuttgart Mu-
seum, formerly the Naturalien Cabinet.
Records—SOUTH AFRICA: Cape Peninsula to Port St. John’s
(Koch, 1949, p. 509); Cape Point (AWBP coll.); Port Alfred
(AWBP coll.); Buluga Bay, East London (Mrs. N. Prior).
Patella exusta Reeve, 1854
(Pl. 9S, tig. 1; Pl. 104, fig. 3)
Range— Mauritius.
Remarks—The typical subspecies seems to be
confined to Mauritius, although the wider-rang-
ing subspecies pica Reeve, sometimes occurs
along with it. The distinctive character of exusta
is the dull-black, thick outer layer of the shell,
which also produces the narrow, black internal
border. It is unfortunate that the better known
name, chitonoides has to fall as a synonym of
exusta.
Description—Shell of moderate size, up to54 mm.
(2% inches) in length, solid, rather depressed, elon-
gate-ovate, and decidedly narrowed anteriorly.
Apex subcentral to about the anterior third.
Sculpture consisting of about 20 narrow sharply
raised primary radial ribs, and 3 or 4 weak radial
threads in the wider interspaces, over the post-
erior half of the shell. The margin is broadly and
shallowly corrugated. Colour of exterior, when
not encrusted or abraded, uniformly dull black;
interior pale bluish to pinkish white, with a nar-
row black margin. The spatula is weakly defined,
and sometimes partly clouded by a pale fawn
callus.
[01-653]
128 Subgenus Scutellastra
Measurements (mm. )—
length width height
54.0 38.0 13.0 syntype of chitonoides
53.0 44.0 14.0 lectotype of exusta
48.0) 38.0 16.0 Mauritius
47.0 37.75 11.5. lectotype of chitonoides
45.0 35.0 12.5. Mauritius
Synonymy—
1854 Patella exusta Reeve, Conch. Iconica, pl. 15, figs. 35a, b.
(locality unknown).
1854 Patella chitonoides Reeve, Conch. Iconica, pl. 21, figs. 52a,
b. (locality unknown). Dec. 1554.
1891 Patella (Scutellastra) exusta Reeve, Pilsbry, Man. Conch.
vol. 13, p. 98, pl. 24, figs. 9, 10.
1891 Patella (Scutellastra) pica form chitonoides Reeve, Pilsbry,
Man. Conch., vol. 13, p. 98, pl. 26, figs. 28, 29.
Types—The types of both exusta and of chito-
noides are in the British Museum (Natural History).
That of exusta consists of three syntypes glued to a
tablet. One of these, measuring 53 x 44 x 14 mm.,
matches Reeve’s figure, pl. 15, fig. 35, and is here
nominated lectotype. The other two specimens on
the tablet are the Californian Acmaea pelta Esch-
scholtz, and may have been added later than Reeve.
The type series of chitonoides also consists of three
syntypes, and the one matching Reeve’s fig. 52,
Plate 98. Fig. 1. Patella (Scutellastra) exusta Reeve, 1554.
Mauritius, 48 mm., AWBP. coll. 628. Figs. 2-4 Patella
9
(Scutellastra) exusta subspecies pica Reeve, 1854. Fig. 2.
Mauritius, 38 mm., AWBP. coll. 627. Figs. 3.4. Mahé,
Seychelles, 42-45 min., AWBP coll. 46135.
A. W. B. Powell
Patellidae
measuring 47 x 37.75 x 11.5 mm. is here nomi-
nated lectotype.
Records—Locality unknown for the types of both exusta
and chitonoides. MAURITIUS: (AWBP coll.); Vacoas Point,
3 miles south of Mahebourg (Ruth Ostheimer and Virginia
Orr, Nov. 20, 1960).
Patella exusta
subspecies pica Reeve, 1854
(Pl. 98, figs. 2-4)
Range—Islands of the Indian Ocean, Mauritius to
Seychelles.
Description—Shell of similar size and shape to
exusta typical, being elongated and noticeably nar-
rowed anteriorly, but the coloration is different, for
instead of the whole of the outer surface being
black, that colour is confined to radiate lines or
streaks, often short and intermittent, and present
only in the primary rib interstices, the rest of the
exterior of the shell being white. Internally the shell
is porcellanous-white with the spatula irregularly
stained orange-brown, and at the margin the dark
external pattern shows through, forming an inter-
mittent narrow border.
Measurements (mm.)—
length width height
49.5 38.00 14.0 — lectotype
47.0 37.25 10.0 Mauritius
44.5 36.25 10.5 Mahe, Seychelles
38.0 30.00 6.5. Mauritius
30.4 26.00 5.0 Seychelles
Synonymy—
1854 Patella pica Reeve, Conch. Iconica, pl. 19, figs. 45a-c.
1891 Patella (Scutellastra) pica Reeve, Pilsbry, Man. Conch.,
vol. 13, p. 97, pl. 22, figs. 9, 10, 13, 14; pl. 59, figs. 47-
49 (not pl. 26, figs. 28, 29).
1863 Patella moreli Deshayes, Cat. Moll. Réunion, p. 43, pl. 6,
fig. 13.
1863 Patella levata Deshayes, Cat. Moll. Réunion, p. 44, pl. 6,
fig. 14.
Records—*‘South Seas” (type); MAURITIUS: (AWBP coll.);
Vacoas Point, 3 miles south of Mahebourg (Ruth Ostheimer and
Virginia Orr, Nov. 20, 1960). SEYCHELLES: (AWBP coll.);
Mahé (AWBP coll.); Frigate Island (AWBP coll.). REUNION:
(Deshayes, 1863).
Types—The type series of pica is in the British
Museum (Natural History) and consists of three syn-
types mounted upon a tablet. The one measuring
49.5 x 38 x 14 mm. is here nominated lectotype.
[01-654]
November 27, 1973
Patella flexuosa Quoy and Gaimard, 1834
(Pl. 65, figs. 8, 9; Pls. 99, 104)
Range—Andaman Islands and the tropical Pa-
cific as far east as the Tuamotu Islands.
Remarks—This exceedingly variable species has
had many names, and unfortunately the one
under which it has usually appeared, stellaeformis
Reeve, 1842, must fall as a synonym of the earlier
flexuosa Quoy and Gaimard, 1534.
The species is intertidal and a shallow-water
dweller, living attached to coral rock, or some-
times upon the outer and inner surfaces of large
shells. Examples living attached to shells tend to
be of lighter build, and of more circular outline
INDO-PACIFIC MOLLUSCA, vol. 3, no. 15
Patella = 129
than those from coral-rock faces, but that is not
invariably so, and all manner of shapes, sculpture
and coloration is encountered, irrespective of
station.
The species varies between 14 mm. and 41
mm., is circular to elongate-ovate in shape, may
be almost flat to moderately elevated, thin or
solid. The strength of the external ribbing deter-
mines the degree to which the margin is scal-
loped. The exterior is dull-white, sometimes spar-
ingly speckled, lined, or with zigzag dark-brown
markings in the rib interstices, but the outer
surface is usually lime-encrusted. The interior is
porcellanous, more or less white, and the spatula
Plate 99. Figs. 1-9. Patella (Scutellastra) flexuosa Quoy &
Gaimard, 1834. Fig. 1. Rouw Island, Aoeri Islands, West
New Guinea (stellaeformis form), 30 mm., AWBP. coll. 39227.
Figs. 2, 3. Michaelmas Cay, off Cairns, North Queensland,
29-34 mm. (topotypes of inquisitor), AWBP coll. 46062.
Fig. 4. New Caledonia, 31 mm. (topotype of intraurea),
AWBP. coll. 45672. Figs. 5, 6. Wake Island, 31-34 mm.,
AWBP. coll. 204725. Figs. 7, 8. Paea, Tahiti, Society Islands
33-35 mm. (paumotensis form), AWBP coll. 115. Fig. 9.
Bikini Island, Marshall Islands, 35 min., AWBP coll. 48226
[01-655]
130. Subgenus Scutellastra A. W
may be white, yellowish, orange-brown, or some-
times dark-slate.
Description—Shell rather small, up to 42 mm.
(1% inches) in length, typically roundly-octagonal,
with the apex nearly central, sculptured with 8 or
9 rounded radial folds that project strongly at the
margin. The entire surface, folds and interstices
alike, is densely sculptured with secondary, crisp,
scabrous to spinose cords. In some forms the pri-
mary and secondary radials become nearly equal,
in which cases the octagonal outline is modified
to a crenulated oval. Colour as described under
remarks.
Radula—Formula 3 + ] + (2+1+2) + 1 + 3, very
similar to that of exusta pica and kermadecensis.
Prashad and Rao figured the radula of their
‘tara, which compares closely with the radula of
a Cook Islands specimen. Prashad and Rao con-
sidered their species to be related to the South
African granularis, but in that species the cusps
are better formed, parrot-beaklike, and the shell
is quite unlike the flexuosa group in form, size,
texture and coloration.
Measurements (mm.)—
length width height
41.0 35.0 17.5 Fakarawa, Tuamotus
39.0 30.0 9.5 ‘Tokorava, Tuamotus
33.0 30.0 8.0 Michaelmas Cay (type of
inquisitor)
30.0 28.0 9.0